file
Paul Gallagher
filename="Book_proposal.txt"
THE ENTANGLED BANK OF EVOLUTION
DARWIN AND BEYOND
Man ... will never die, because there may never be a taxonomical point
in
his evolutionary progress that could be determined as the last stage of
man
in the cline turning him into Neohomo, or some horrible throbbing slime.
-Vladimir Nabokov, Ada, p. 406.
The NEXA Program for Science-Humanities Convergence at San Francisco
State University, which was founded by a development Grant from NEH in
1975, is planning an interdisciplinary volume entitled, "The Entangled
Bank of Evolution: Darwin and Beyond." Principle Investigator: Michael
S. Gregory, Director, NEXA Program, SFSU; Co-Investigators: Michael T.
Ghiselin, Chair, the Department of the History and Philosophy of
Science, California Academy of Sciences; Paul C. Gallagher, Associate
Editor for Research and Planning, H-NEXA, the Science-Humanities
Convergence Forum (H-Net/MSU/NEH).
No intellectual matter of our time is richer in complexity than the
subject of evolution, which necessarily involves questions about the
origins and nature of life itself, the various and competing modes of
explanation for development in general and human development in
particular, and the implicit questions of human nature and human
destiny.=20
We believe that this book is a fitting way of looking, as we enter the
new millennium, at such matters as predetermination of behavior
(sociobiology), of thought (evolutionary psychology) and even ethics:
E.O. Wilson's "ethics of the gene," as opposed to rational freedom,
human rather than gene-fixed ethical behavior, and developmental
indeterminacy. We are involving notable specialists in the biological
and other sciences to join philosophers and others humanists in this
comprehensive discussion of human origins, human nature, and human
potential. We are asking humanists and scientists to combine the
perspectives of their two great domains of knowledge in the kind of
cooperation pioneered by NEXA:
A Program for Convergence. In this survey of evolution as idea, as
hypothesis, as theory, and now all but law, we intend to assess such
areas as the following:
I Origins of Life -On Earth and in The Cosmos
II. Evolution - Theory and Concept, Past and Present
"Evolution" and "Progress" - the History of Social Ideas
IV. Evolution Beyond Biology - Evolutionary Applications in Other
Sciences
V. Evolution Among The Primates - Our Hominid Origins and Our
Ancestors
VI. Entities of Evolution-Species and Beyond
VII. Rates and Kinds of Evolutionary Change -- Slow
Accumulation, Sudden Leaps, or Neutral Drifts
VIII. Environmental Transactions - Transducers and Transponders in
Biochemistry
Sociobiology and Evolutionary Psychology - "The Ethics of the Gene."
Michael S. Gregory
Director, NEXA: a Program for Convergence
San Francisco State University
-------------------------------------------------------------------------=
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SECTION C: PROJECT DESCRIPTION
Introduction.
"Evolution After Darwin," November, 1959
"The Entangled Bank of Evolution: Darwin and Beyond" is designed to be
the first full-scale inter-disciplinary assessment of the Matter of
Evolution since the 1959 centennial symposium, "Evolution After Darwin,"
held at the University of Chicago on the anniversary of the publication
of Charles Darwin's On The Origin of Species. That event involved 44
eminent scholars from 10 nations representing seven disciplines and
three great divisions of knowledge: inorganic sciences, life sciences,
and human sciences, speaking in five panels:
Panel I: THE ORGINS OF LIFE
Panel II: THE EVOLUTION OF LIFE
Panel III: MAN AS AN ORGANISM
Panel IV: THE EVOLUTION OF MIND
Panel V: SOCIAL AND CULTURAL EVOLUTION
.
Each panel had two co-chairs, and had from seven to nine panelists.
The monumental publication, Evolution After Darwin, edited by Sol Tax
(Chicago; 1960) that eventuated from this event was divided into three
volumes of essays: "The Evolution of Life," "The Evolution of Man, and
"Issues in Evolution," the text devoted to the panel presentations.
"The Entangled Bank of Evolution - Darwin and Beyond." Prologue.
Since 1959, much has changed and developed in matters concerning
evolution. It is very largely a
new world.
On the one hand, we now have in prospect the discovery of the mechanisms
by which life itself
appeared, and dead matter in the kingdom of entropy and ultimate
disorder became, under still-unknown
conditions, suddenly alive, pulsating and self-replicating. For the
first time in the history of the our world, the "anti-entropic"
phenomenon of Life emerged from what John Milton called "The Realm of
Chaos and Old Night." Additionally, we are now learning more about what
conditions on other planets are likely to be conducive to life, and new
probes to Mars and other worlds are seeking clues and perhaps evidence
of life evolving elsewhere in our solar system, and eventually in our
galaxy. Whether that evolution will follow the Earth pattern or
represent divergent developments in divergent conditions is a fit
question to be considered as we cross the border into the Third
Millennium.
On the other hand, in the past few years new vistas of genetic
predetermination have opened up -predetermination of behavior and mind,
and the motivating entity has devolved to the level of the gene. Where
before one might speak of species (and to some extent the individual) as
the unit of evolutionary process, we are now persuaded to accept the
gene in its entwining chromosome as the
unit of selection, and by implication the ontological locus of life
itself. In this view bodies, minds and behaviors are no more than the
temporary and renewable expressions and instruments of the ineluctable
will of the gene, and the gene itself no more than a particular
arrangement of nucleotide bases known as DNA. The will of the gene is
expressed in molecules of ribonucleic acid (RNA) that produce its own
metabolism and proteins that allow it to transport its particular
hereditary features by reproduction and adaptive selection into an
endless future - a process we know as evolution.
Four thousand million years on, what was to be the fate of the ancient
replicators.=20
They did not die out, for they are past masters of the survival arts.
But do not look=20
for them floating loose in the sea; they gave up that cavalier freedom
long ago. Now=20
they swarm in huge colonies, safe inside gigantic lumbering robots,
sealed off from
the outside world, communicating with it by tortuous indirect routes,
manipulating
it by remote control. They are in you and in me; they created us, body
and mind; and
their preservation is the ultimate rationale for our existence. They
have come a long=20
way, those replicators. Now they go by the name of genes, and we are
their survival=20
machines.
- Richard Dawkins, The Selfish Gene
Curiously, in this respect, evolution has followed contrary paths: one
upward to the broad heavens, the other downward to the microscopic level
of biochemistry, where new things are forever brewing out of congeries
of acids (RNA) engaged in variation and selection, in ceaseless
competition with other chemical congeries of similar type. This
divergence in scientific purview and perspective in itself marks a
division, perhaps a diremption, of the modern mind. Whereas for
millennia, mankind followed the cosmological precept, "As above, so
below," we now have the not always comfortable companion precept, "As
below, way below, so above."
One striking feature about modern thought generally is that it tends to
be almost universally reductionistic. Reductionism is without question
the mode of modern biology. The clues to later complexity are believed
to lie at the lowest, most formative, levels of analysis. There is much
to recommend this way of understanding. It gives us a picture of
earliest developmental structures. It in effect allows one to reverse
the process of evolution. It provides a time-machine that can take the
observer back to earliest beginnings of the higher, more developed,
forms under consideration. Here is a bedrock of factual information for
the structural and developmental biologist, enabling a comprehensive
investigation of a finite and delimited set of features. This has the
value of a nether ne plus ultra, and provides indisputable facts without
the intrusion of conjecture. However, in terms of describing,
prescribing or proscribing higher levels of organization, it is flawed
by the fundamental logical error of pars pro toto, although this fact
has yet to receive adequate attention.
Reductionism has found a home in the humanities, as well, in the
movement called Postmodernism. Here is a movement that seeks the basic
elements of human knowledge and experience at the lowest of ontological
and epistemological levels. Postmodernism seeks at the very bottom of
our traditional mode of thinking an ultimate objectivity devoid of
values, preconceptions or even ideas. "Modernism may be seen as an
attempt to reconstruct the world in the absence of God," said Bryan
Appleyard, meaning that the world has no discernible natural order, and
that all presumed order is a cultural construct which must be discarded.
The French semiologist Jean Baudrillard puts in this way: "Postmodernity
is the simultaneity of the destruction of earlier values and their
reconstruction. It is renovation within ruination," where ruination is
traditional "modernist" meaning, and renovation is a pure and utterly
detached nullity. [1] And adding a presiding null status quo ante to all
experience, Jean Fran=87ois Lyotard has said, "A work can become modern
only if it is first postmodern. Postmodernism thus understood is not
modernism at its end but in the nascent state, and this state is
constant." [2] Thus, while there are similarities of process in
biological science and postmodernist humanities, in that they both
distrust higher-level formulations and seek truth in fundamentals, the
truths they find are worlds apart. Biology seeks and finds unadorned
facts; Postmodernism finds a basic void, where everything is both true
and untrue at the same time. As Samuel Beckett put it, "There is no lack
of void." [3]
In terms of our proposal, there is here an apparent agreement about the
locus of meaning discovered by reductionism; just as "modernist"
experience has only spurious meaning, genetic reductionism has
significance but no inherent meaning. Genes do not "mean" and they have
no intention; indeed, to imagine that they do would be to invoke
teleology, which is banished from Darwinist evolutionary thinking. The
sole functions of genes are to prevail and to reproduce. Thus, there is
a surprising univocal quality about the most disparate forms of modern
thought: Meaning is a fossil in the record of discarded human values.
Any number of other terms now can substitute: power; velocity;
self-assertion; dominance. And each of these, not coincidentally, is a
feature of the newest branches of evolutionary theory: Sociobiology and
Evolutionary Psychology. And these fields are both the progenitors and
the offspring of reductionistic methods of analysis:=20
[S]ome sort of dualism is the inevitable consequence of any sort of
reductionist
materialism that does not in the end wish to accept that humans are
'"nothing but"
the motion of their molecules. Dualism was a solution to the paradox of
mechanism that would enable religion and reductionist science to stave
off for
another two centuries their inevitable final contention for ideological
supremacy.
- Richard Lewontin, Steven Rose, Leon Kamin, Not in Our Genes, p.
46.
Oddly enough, the polar opposite of reductionism is not cited, and one
searches vainly for its name among challengers to this system. If monism
or holism are the intended opposites to dualism, we have yet to find
their presence in the reductionistic counterpoise. Logically,
reductionism should find a mating opposite, but that way seems barred by
the metaphysics of theology. Curiously, it was just this sort of impasse
that entangled conceptions of evolutionary rate and time. As Niles
Eldredge puts it:
[U]niformitarianism and its commingled twin, gradualism, have become
nearly
synonymous with reductionism. The notion that what we see around us
today is both
necessary and sufficient to explain the evolutionary histories of the
earth and
of life has become a rationale for probing no further. In some instances
it has
meant insisting that the data and theory of one narrow outlook or
discipline
are all that collectively we will ever need to reach full understanding.
- Niles Eldredge, The Pattern of Evolution, pp.
39-40.
This symposium will not encounter a question more basic and tendentious
that than of reductionistic analysis and its productive but strictly
limited yield of information about evolution and, indeed, the about the
lives and natures of higher organisms. Let us mark this important
impasse with this luminous insight not into levels of analysis, but into
the limits of human cognition itself:
In each age of the earth distinguished by high activity there will be
found at its
culmination, and among the agencies leading to that culmination, some
profound cosmological outlook, implicitly accepted, impressing its own
type
upon the current springs of action. This ultimate cosmology is only
partly
expressed, and the details issue into derivative specialized questions
of violent
controversy. The intellectual strife of an age is mainly concerned with
these
questions of secondary generality which conceal a general agreement upon
first principles almost too obvious to need expression, ad almost too
general
to be capable of expression. In each period there is a general form of
the
forms of thought; and, like the air we breathe, such a form is so
translucent,
and so pervading, and so seemingly necessary, that only by extreme
effort
can we become aware of it.
- Alfred North Whitehead, Adventures of Ideas, 1933
Thus, modern evolutionary biology bears the stamp of our Third
Millennium worldview: it is a divergent worldview, perhaps disturbingly
so. We are looking out with telescopes and probes to the largest context
of our attention - the universe; and we have brought to bear for the
study of evolution on Earth the most focused and delimited
epistemological tool of our attention -reductionism. Between "Above" and
"Below" there is no longer an "As" pointing in either direction, and we
have adopted or had thrust upon us (for now all but forgotten heuristic
reasons) an epistemological splitting of the natural world, a
hierarchical diremption, with for at least the non-scientist, an
accompanying and perfectly reasonable anomie.
Chapter I: ORIGINS OF LIFE - ON EARTH AND IN THE COSMOS
Chapter II: EVOLUTION - THEORY AND CONCEPT, PAST AND PRESENT
Chapter III: "EVOLUTION" AND "PROGRESS" - THE HISTORY OF SOCIAL IDEAS
Chapter IV: EVOLUTION OUTSIDE BIOLOGY - EVOLUTIONARY APPLICATIONS IN
OTHER
SCIENCES
Chapter V: EVOLUTION AMONG THE PRIMATES - OUR HOMINID ORIGINS AND OUR
ANCESTORS
Chapter VI: ENTITIES OF EVOLUTION - SPECIES AND BEYOND
Chapter VII: RATES AND KINDS OF EVOLUTIONARY CHANGE - SLOW ACCUMULATION,
SUDDEN LEAPS, OR NEUTRAL DRIFTS
Chapter VIII: ENVIRONMENTAL TRANSACTIONS - TRANSDUCERS AND TRANSPONDERS
IN
BIOCHEMISTRY
Chapter IX: SOCIOBIOLOGY AND EVOLUTIONARY PSYCHOLOGY - "THE ETHICS OF
THE
GENE"
- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -
- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -
- -=20
Chapter I: ORIGINS OF LIFE - ON EARTH AND IN THE COSMOS
Two of the great mysteries that confront science and have compelled the
imagination are how life came into being from non-living matter, and
whether life, perhaps including intelligent life, exists beyond the
Earth. These questions have inspired some of the most distinguished
scientists and finest essayists; the origin of life is an especially apt
topic, not only because of its intrinsic interest, but because so many
eloquent people are available to discuss current research.=20
A popular theory for the origin of life supposes there was once a "RNA
world." It is often assumed
that prior to the earliest cells, "precellular" life existed, but the
early molecule being replicated was not DNA but RNA. RNA presents a
solution to what came first, genes or enzymes, by being both, and in
turn being able to be transcribed into DNA. A string of Nobel Prize
winners developed the idea: Francis Crick first guessed that RNA might
be the first genetic molecule, catalyzing its own replication; Thomas
Cech proved that RNA can act as an enzyme; and Walter Gilbert coined the
term, "RNA world," wherein RNA molecules catalyzed the assembly of other
RNA molecules from free nucleotides. Then, through mutation and
selection, these molecules developed RNA adaptor molecules to bind amino
acids and arranged these amino acids along a RNA template to create the
first proteins. When DNA finally appeared, the template in the RNA
molecule was transferred to DNA by reverse transcription, and RNA took
on the role of intermediate between DNA and proteins. Later research has
shown that complex functional RNA ligases can form by chance alone.
However, the RNA world is not accepted by everyone. RNA may be too
fragile and difficult to synthesize. Boehler, Nielsen and Orgel propose
that prior to the RNA world there was a PNA world based on peptide
nucleic acid. Some believe bare DNA came first; others think that
proteins came first; Freeman Dyson thinks they both arose separately. A.
G. Cairns-Smith has proposed the "Clay Life" theory, in which clay
crystals formed a template for the early genome. Harold Morowitz and
David Deamer think that proto-cells came first, before genes. Jeffrey
Bada thinks the early Earth was cold, and life originated in cold water
under the ice. Claudia Huber and Guenter Waechtershaeuser think life
originated near undersea volcanic vents, where the heat and minerals
could favor necessary reactions. Thomas Gold thinks life might have
originated in the Earth's crust. There has been great recent interest in
"extremophiles," microbes that live in ice, in the Earth's crust, or
near volcanic vents, dependent on sulfur and iron for energy, in part
because of their possible resemblance to the earliest life and to life
that may exist on other planets. The tiny bacteria-like fossils some
perceive in the Martian meteorite, ALH84001, may have found an earthly
parallel in nanobacteria, the smallest bacteria, which may or may not
form most of the Earth's biomass and which may or may not control much
of the earth's surface chemistry, in addition to being a likely cause of
kidney stones.
Manfred Eigen emphasize the role of self-organization in the origin of
early life. Eigen, in Steps Towards Life : A Perspective on Evolution
and The Hypercycle : A Principle of Natural Self Organization,
speculated on the role of hypercycles in life's origin. Hypercycles are
a kind of reaction network based on "nonlinear autocatalysis"
(self-replicating molecules linked by cycles of catalysis) that arises
from a molecular population and evolves toward higher complexity. Stuart
Kauffman, in the Origins of Order, also argues that complex assemblages
of chemicals can have self-organizing properties. Once life appeared it
quickly became cellular. The earliest sedimentary rocks on earth contain
cells, and evidence of cellular metabolism begins at least 3.8 billion
years ago. The discovery of some of the earth's earliest fossils is
recounted in Cradle of Life : The Discovery of Earth's Earliest Fossils
by J. William Schopf. The subsequent three billion years of evolution,
once almost unknown, have been the subject of considerable study over
the past decades. Some of the major events of the Archean eon are being
explored -- the origin of photosynthesis, the origin of the earth's
oxygen atmosphere, the origin of the eukaryotes, the origins of sexual
reproduction, as described , for example, in Lynn Margulis and Dorion
Sagan's Microcosmos : Four Billion Years of Evolution from Our Microbial
Ancestors.=20
Theories of the origins of life are numerous , but we might distinguish
between those who think life is almost inevitable given a certain range
of prebiotic conditions and those who think the origin of life unlikely.
These theories are reflected in the search for life elsewhere in our
solar system: on Mars; on the moons of Jupiter, Europa and Callisto; and
on Saturn's moon, Titan. It is now a common point of view, defended by
Christian de Duve and many others, that life originated on Earth in a
matter of centuries, or in an even briefer span, and that life is
probable on any world where there is liquid water. Hence the liquid
water suspected to lurk beneath the surface of Mars or the oceans
thought to exist on Europa and Callisto may harbor life, waiting to be
found. This search received attention when it seemed to reach a
successful conclusion when traces Martian life might have been found in
1996 in a meteorite of Martian origin.
Beyond our solar system the search for radio signals from intelligent
life continues. The probability that there are technological
civilizations nearby in our galaxy, or whether intelligent life on other
worlds would resemble humans, are questions often debated among
biologists, since it is not clear that the historical contingencies that
led to human evolution should have parallels on other worlds.=20
The discovery of life on other planets opens up the prospect of a
universal biology. Life elsewhere would allow for the first time life on
Earth to be placed in comparative perspective, the better to understand
what is necessary and contingent in living things.
Chapter II: EVOLUTION - THEORY AND CONCEPT, PAST AND PRESENT
Evolutionary theory is almost entirely a product of the nineteenth and
twentieth centuries. At the beginning of the twenty-first it seems most
appropriate that we reassess our ideas about where it came from, and
where it seems to be headed in the present.
The received historical account of the history of evolution clearly
distinguishes between claims that evolution has in fact occurred on the
one hand, and what causes it on the other. Darwin gets credit mainly for
proposing a plausible mechanism, natural selection, but his arguments
for evolution as a fact seem to have been much more convincing than
those of his predecessors. At any rate it has seemed ironic that Darwin
was able to get evolution, but not its mechanism, accepted by the
scientific community as a whole.
Supposedly, there were serious problems with his theory, largely due to
the lack of a proper theory of heredity. But religious, ideological and
philosophical objections have been invoked as well. Efforts to champion
alternatives to natural selection, such as Lamarckism, Geoffroyism,
mutationism, orthogenesis, and saltationism persist to the present day.
According to the traditional historiography, such alternatives became
untenable due to the rise of modern genetics and the emergence of a
Synthetic Theory in the 30s and 40s. And yet the theory that emerged was
very much like the one that Darwin himself had proposed.=20
Some authors have seriously questioned whether there really was a
synthesis and what actually happened continues to be debated by
historians. Furthermore there have been numerous and vocal claims that
the theory is in the process of collapse. Indeed it is sometimes treated
as a social construction, an orthodoxy, perhaps a myth.
Placing the debates in historical retrospect should help at least to
clarify some of the issues and lead to some productive discussion. This
module will give an overview of the history of evolutionary biology,
with two main goals in mind. First it should help our audience better to
appreciate the issues. Second, it should contribute to our understanding
of the received view, and its alternatives. How viable are various
revisionist efforts apt to be?=20
Chapter III: "EVOLUTION" AND "PROGRESS" - THE HISTORY OF SOCIAL IDEAS
It perhaps seems inevitable today to imagine that "evolution" and
"progress" have always been synonyms, but that is not the case. Indeed,
the word "progress" in its modern sense appears to be all but unknown
until the European seventeenth century, and its true origin remains
obscure. Cross-culturally, there are simulacra of this idea but no real
equivalents (e.g., time configured as cycles, gyres, etc.) Time
configured as rectilinear and inclining upward is a distinctly modern
and western conception. Confucianism, one of the half dozen major and
enduring worldviews, configured time precisely as the obverse:
rectilinear and, from the Golden Age of Yao and Shun, inclining ever
downward. The measure of time is inevitably associated with a locus of
value: past (Confucian and other strictly conservative worldviews);
present (Navajo and Hopi, where all time is time-present), and future
(the modern, western worldview as the most distinctive example in the
human record.=20
"Evolution" up until about 1830 meant development - but individual
development only. One spoke of a man's or woman's evolution, from cradle
to grave. The modern novel in one of its commonest forms, the
Bildungsroman, or biographical novel, means literally "novel of
development," and Kunstlersroman considered the (always romantic)
"development of the artist."=20
The exact point at which "evolution" assumed its modern form is obscure,
but Lamarck and Erasmus Darwin (Charles Darwin's grandfather) wrote a
treatise about evolution called Zoonomia (1794-96), which employed the
term well before it came into popular usage, and probably inspired the
work of the great invertebrate zoologist (le Chevalier de) Jean Baptiste
Pierre Antoine de Monet Lamarck (who kept his petit-noble head while all
around were losing theirs), the Zoological Philosophy (1809). From that
point forward, "evolution" became bonded to species, and it would seem
strange these days to hear of the evolution of a person, except perhaps
ironically.=20
But in terms development, the notion of "progress" had been for
centuries hovering near. Since this matter is highly detailed, it will
have to suffice to present the stages through which is passed in tabular
form, thus:
Entelechy. Classical Greece. The movement of a thing from a baser to a
higher form. All things had entelechy (hence the Medieval alchemist's
goal of creating gold from lead was based upon a perceived scientific
principle that needed only magical encouragement; even Newton was not
free of magical thinking). For Artistole, entelechy meant the realized
essence of a thing. Elsewhere in Greek philosophy, it meant the force
driving toward that realization (the ultimate source of Lamarck's
sentiment interieur).
Fixity of Species. Judeo-Christian Europe. The species were created
during the period of the Genesis and remain unchanged both in form and
number. No evolution. No new forms. No extinctions.
The Great Chain of Being. Scala Naturae, scale in the musical sense.
Later hybrid of Fixity of Species, imparting unchanging hierarchy,
ordained by God, from the lowest (worm), through the various animal
forms to the very top, Himself. Later to become a ready tool of colonial
apologia.
Serial Creation. The doctrine of Fixity losing ground to paleontology.
Fossils, first thought to be "thunderstones," or Devil's work to deceive
mankind into unbelief, finally had to be reconciled with Scripture: the
beginning of a long retreat. This doctrine had God for reasons best
known to Himself, returned to Earth periodically to erase a page of His
Genesis and introduce another round of Creation. For means, He adopts -
Catastrophism. The risky business of putting the Creator in charge of
massive misfortune, and indeed extinction. This doctrine explained the
Jobian question of the Heavenly origin of things men call on Earth call
evil, and also reconciled the physical evidence of fossils and the
stratigraphic record with Divine Will. Catastrophism was God's eraser,
clearing the slate for the next round of Serial Creation.
Progressivism. Here begins the great conflation between Change and
Improvement, in both of two senses. First, it gave God a reason for His
Serial Creations by Catastrophic means: he just wanted to make things
better each time. Thus, the Almighty begins to look like Hamlet's
apprentice, never getting it right and having to try it over and over
again. Second, it re-introduced the powerful notion expressed in the
simple imperative first observed by Heraclitus: "All things change," and
added the distinctly modern secondary imperative: "All things change for
the better." From God as static perfection to God as ever-altering
Improvement.
Uniformitarianism. Change having finally been forcibly admitted into
God's plan, the question of kind and rate became foremost (see
description below of Chapter VII: Rates and Kinds of Evolutionary
Change. The Uniformitarian view, which now prevails and is in accord
with the predominant scientific mode of reductionism, holds (contra
Catastrophism) that the physical evolution of the Earth requires no
Catastrophic changes to comprehend the state of the world we see
presently before us. The forces observable at work on the planet today
are the same forces that have always prevailed, and the Creator, if any,
has put his Creation on cruise-control by agency of secondary causes,
and His direct work as First Cause is nowhere to be seen (and there is
no need to suppose it was ever present in the world). In terms of
geomorphology, Sir. Charles Lyell was the foremost exponent of
Uniformitarianism, and was held in almost religious awe by Darwin;
Darwin received one volume of Lyell's monumental work, Principles of
Geology (1830-1833), in Argentina during the voyage of the Beagle.
Gradualism. A logical, but unnecessary, correlate of Uniformitarianism.
There is nothing in the doctrine of Uniformitarianism that constrains
naturalistic changes to be slow or even steady. The question of rate and
kind of evolutionary change is one of the two unifying features of this
symposium; the other is cause of change. Nowhere is The Bank of
Evolution more Entangled than on these questions. Darwin probably
unnecessarily conflated uniformity with slowness, and it later caused
him much pain. When Lord Kelvin, calculating the age of the Earth on the
basis of how long the Sun had been burning (like a lump of coal), denied
to Darwin the enormous sweep of geological time he required for his
theory of evolution, Darwin wrote to a friend, "I take the Sun much to
heart," upon receiving Kelvin's unsettling news.=20
It is difficult to say whether Progress was the tree and Evolution the
ivy that grew up that tree, or the other way around, but is seems clear
that Improvement was the seed from which both sprang. The very notion of
passage through time producing a building-betterment, Bildung is, as
noted, a peculiar and new one on the world scene. Very few people prior
to the modern, perhaps the industrial-modern, era have believed it. [1]
Many forces have conspired to reify and legitimize this entirely
metaphysical notion. It has taken on myriad interlocking self- and
cross-validating forms, now so entirely doctrinal in their authority
that few would attempt to question the premise from which they derive:
for example, Ricardo's [2] seminal argument for endless betterment
through time, and Mill's championing competition of ideas as the only
way to truth [3].
Even ideas in opposition to the competition-progress mode hurled
cobblestones picked up from the same street (an exact metaphorical
expression of Whitehead's warring "forms of thought" overlying a
basically uniting "general form of the forms of thought," supra.
Saint-Simon [4] had forecast an ideal technocratic State, but one which
must be governed in such a way as to control the vagrancy's of unruly
human nature. Robert Owen [5] saw Utopias of regulated cooperation.
Bakunin [6], for the first time entirely flying the coop of control and
regulation, saw Improvement imprisoned by an evil opposite force, any
form of government, let or hindrance, on naturally human goodness. And
Marx [7] foresaw the triumph of human goodwill and cooperation after
competition, an essentially alien and malevolent historical-industrial
accident, had exhausted itself in economic Armageddon.
Taking Mill again as example, we find Malthusian economic
competitiveness just beneath the surface
of a benign Utilitarianism:=20
As for charity, it is a matter in which the immediate effect on the
persons directly=20
concerned, and the ultimate consequence to the general good, are apt to
be at=20
complete war with one another. [8]
The belief in Improvement has in the past 200 years become tempered by
caveats hinting of special circumstances and fragility, especially from
the zealots who love progress and evolution not wisely, but too well.
Thomas Robert Malthus' monumental On Population [9], which profoundly
shaped the thinking of Ricardo, Darwin and Wallace, is as much concerned
with political turmoil from over-believers as he is with proving that
unchecked reproduction will outstrip food supply. The first few pages
are a skilled polemic designed to keep that contagion of political
enthusiasm, the French Revolution, from spreading from the Continent to
an England still bruised by the excesses of the Puritan Revolt. Malthus
rhetorically poses emotion against reason. His argument from population
pits ideas plausibly manifested by "the defender of establishments" and
"the advocate for the present order of things" against the perhaps
well-meaning, but dangerously irrational, "advocate for the
perfectibility of man, and of society." (This is an opposition of ideas
Herman Melville renders in precise symbolic equivalence in the novel
Billy Budd: Foretopman [10].) Malthus truly calls this an "unamicable
contest," which indeed involves the fate of nations.
The great argument with which this symposium is concerned, pars in toto,
is the status of evolution as a science, approached through biology,
zoology, paleontology and biochemistry, and the facts that they discover
and produce, as over against the values, implicit and explicit, which
inevitably color, contextualize and perhaps even in some ways
precondition these discoveries. Like all human endeavors, the study of
evolution brings into play extra-biological phenomena - human ideas,
traditions, hopes, fears, prejudices - as expressed in philosophy, the
social sciences, literature and politics. This event is to be, according
to one's own understanding and predilections, an Omnium-gatherum, a
conclave on Laputa, or a Symposium in the fully Platonic sense.
Chapter IV: EVOLUTION OUTSIDE BIOLOGY - EVOLUTIONARY APPLICATIONS IN
OTHER SCIENCES
It has long been recognized that not only populations of organisms
evolve, but all sorts of other things, notably languages, economies,
science and technology. Although much of the comparison has been weakly
analogical, or little more than metaphor, there are several areas in
which the extension of evolutionary thinking beyond its traditional
limits is receiving enthusiastic support. On the other hand there has
been much discussion about the role of intentionality and
purposefulness, whether Lamarckian mechanisms do or do not apply, and
all sorts of other challenging problems. In this module we consider some
examples that are under active investigation.
EVOLUTIONARY EPISTEMOLOGY has had a profound impact upon philosophy, and
is often associated with the name of the philosopher Karl Popper. Its
main advocate, however, was the psychologist Donald Campbell, who
treated scientific and organic evolution alike as based upon "blind
variation and selective retention." There has been much discussion about
such questions as whether there are any analogues with species or genes.
The time seems ripe for a retrospect of whether such efforts have or
have not been successful. We are asking the philosopher Jagdish
Hattiangadi to give discuss this topic. [Other good candidates:
NicholasThompson, Lisa Lloyd.]
MECHANISMS OF INNOVATION or of CREATIVITY IN THE BROADEST SENSE have
some noteworthy similarities in organic evolution and in the creation of
knowledge and works of art. The
problem of how new organs can come into being through natural selection
was addressed by Darwin in rebutting his critics, and has been
repeatedly been addressed by Darwinians, notably Anton Dohrn (1874,
translation in Ghiselin, 1993) and Ernst Mayr (1960P. The origin of
novelty through change of function is but one of many recognized
mechanisms of broad applicability. Looser analogies and metaphors such
as gene-flow and hybridization and cross-fertilization between
disciplines can be presented as something more concrete and rigorous
when linked to particular cases. Such evolutionary mechanisms can be
opposed to models of discovery that resemble special creation or
saltationism. The time seems ripe for a reconsideration of such topics,
especially the role of analogy.=20
TECHNOLOGICAL EVOLUTION is rich in analogies with organic evolution, for
example in the
striking resemblances between eyes and cameras. It is often invoked as
illustrative of the phenomenon of progress, a topic that has become very
controversial during the last few years and about which the last word
has obviously not been said. The extent to which deliberate planning and
foresight come into play in the changes that we see in technology are
highly debatable.
LEGAL SYSTEMS change in part by legislation, but the common law has been
treated as evolving by a kind of trial and error process in which
litigants are selective agents (Goodman, 1978). There is an active "law
and economics" movement with its own journals and professional
organizations.=20
ECONOMICS AND BIOLOGY are being integrated in several ways.=20
BIOECONOMICS is a new discipline that treats the unity of biology and
economics as an extension of Darwin's Malthusian insights and emphasizes
the identity of their fundamental laws and principles.
EVOLUTIONARY ECONOMICS has somewhat different historical roots and makes
a treats the connection with biology as somewhat more abstract, but sees
something like natural selection operating among firms.=20
Chapter V: EVOLUTION AMONG THE PRIMATES - OUR HOMINID ORIGINS AND OUR
ANCESTORS
Like the origin of life, the origin of humans has occupied a central
place in religion, myth, ideology, and in the self-definition of human
beings. And like the origin of life, the evolution of the human species
poses similar problems to those present in the study of the origin of
life: a shortage of data, a
lack of comparative material, and an abundance of speculation. Both
human uniqueness and the similarity of humans to other animals have long
been recognized - "What a piece of work is a man!... the paragon of
animals!" -but evolutionary science showed that the affinity between
humans and other animals, and indeed every living thing, is based on
genealogy, and Darwin showed that the processes that gave rise to humans
can be entirely natural and without overall guiding purpose. It has been
argued that Darwin's most revolutionary achievement was to topple
mankind's place at the crown of creation. Others disagree: man has
maintained his place atop the great chain of being by hook or by crook.
No less a figure than Alfred Russel Wallace brought divine intervention
back into evolution to account for human beings.
The phylogeny of the human species and its relatives is in dispute. But
as difficult as the problem of phylogeny is, still more tasking is to
explain the evolution of bipedalism, tool-making, gathering, learning,
intelligence, language, consciousness, culture, and other traits that
are, if not unique to humans, uniquely well developed in them. The
difficulties are considerable -- "speech does not fossilize," says John
Shea of SUNY Stony Brook -- but developmental psychologists, cognitive
scientists, linguists, anthropologists, paleontologists, not to mention
sociobiologists and evolutionary psychologists, have gone into the
breach.
The cast of characters is famous: Homo sapiens and its closest living
relatives - chimpanzees, gorillas, and orangutans - and its extinct
cousins: Sivapithecus, Ramapithecus, Aridipithecus ramidus,
Australopithethicus anamensis, A. afarensis, A. africanus, Paranthropus,
Homo rudolfensis, H. habilis, H. ergaster, H. erectus, H.
heidelbergensis, H. neanderthalensis. The australopithecines represent
several species that inhabited a wetter, forested Africa and include
such celebrities as the A. afarensis named Lucy, the "killer apes" of
Ardrey's "African Genesis," and the recently discovered A. garhi. A.
garhi, discovered this year and perhaps a better candidate than A.
africanus for the ancestor of humans, used stone tools to butcher the
bones of animals it scavenged. Climate change accompanied the rise of
Homo habilis and H. erectus, as Africa became drier and savannah
replaced forest. "You go into this period with, in essence, bipedal
big-toothed chimps and come out with meat-eating, large-brained
hominids," says Tim White of Berkeley. H. erectus spread through the
world during the Pleistocene, from England to Java. H. erectus developed
more sophisticated tools: stone hand axes were first. Yet more
sophisticated tools appeared in the middle Paleolithic around 200,000
years ago. Ian Tattersall considers this especially important because it
was not associated with the rise of a new species; instead, it seems to
have been the result of cultural innovation. Some suppose H. erectus
gave rise to modern humans -- the 'multiregional hypothesis' -- but this
is not the majority view, and seems to be contradicted by molecular
evidence. Instead, a separate lineage, Homo sapiens, originated in
Africa and spread throughout the world, perhaps less than 100,000 years
ago, displacing all other human species. H. neanderthalis survived until
at least 33,000 years ago, and H. erectus may have survived until at
least 27,000 years ago. The discovery of 27,000 year old H. erectus or
the discovery of A. gahri (whose specific name means "surprise") are
just a few examples of exciting new discoveries in human evolution:
discoveries that can upset reasonably well-accepted theories.
Human beings are especially distinguished by their mental abilities, and
there has been much speculation on the "prehistory on mind," in which
evidence from the study of the mental development of infants, of brain
structures, of language behavior, and of the archaeological remains of
the tools and other artifacts of early hominids, are brought together to
try to reconstruct the evolutionary history of the mind. Merlin Donald
and Steven Mithen have developed widely read speculations on the
evolution of the human mind. Donald argued there were three great
transitions in The Origins of the Modern Mind. Hominids originally
acquired the ability to imitate older members of the same group with
their bodies. Then they developed the neuronal systems that permitted
them to speak and to tell stories. In the third stage they invented
symbolic systems that allowed them to develop art and science. Mithen
built on Donald's work, added the concept of the modular mind, and drew
parallels between the development of individual humans and the evolution
of humans. The mind of infants is like that of early hominids, with a
rough general intelligence. Later hominids and young children have
specialized intelligences, including one for technology (tools), one for
natural history, one for language, and for social understanding. But
these intelligences are independent; they do not work well together. The
earliest modern humans developed language, but it was closely tied to
social intelligence. It is only in the mature adult, or in the modern
humans of the past 30,000 years, that a
general intelligence emerged for the purpose of "meta-representation,"
allowing humans to join
together the separate intelligences to solve problems and create human
culture. The Chapter will likely address the evolution of the capacity
to use language, a topic that has received great attention recently. It
is usually supposed that the ability to use language is recent; that
only anatomically modern humans are capable of speech. The earliest
modern humans of 150,000 years ago are the subject of controversy. They
seem almost identical to modern humans and should be capable of speech.
Their artifacts are impressive -- burials, trade, the beginnings of art,
and sophisticated tools -- but these do may not require verbal language.
The artifacts of modern humans did not differ too greatly from those of
the speechless Neanderthals with whom they coexisted in the Middle East.
However, by the upper Paleolithic of 40,000 years ago, highly
standardized and highly specialized tools, seafaring, and sophisticated
cave painting all indicate that the origin of language must have
occurred, both for transmitting information and sharing social meanings.
Whether earlier humans could speak is debated, and indeed some argue
that earlier hominids - even autralopithicines - possessed language.
Researchers commonly take positions on what the chief purposes are that
language serves, and take a stance on Chomsky's universal grammar.
Terrence Deacon, for example, offers a theory of the origin of language,
emphasizing the separateness of language and intelligence, and proposes
adaptive scenarios for the evolution of the ability to learn and process
symbolic references. He rejects alternative theories in which greater
intelligence, articulatory abilities, or grammatical abilities were
prerequisites for the evolution of language. In fact, symbolic
references may not have been originally verbal, Deacon thinks, but
instead tied into the ritual of social contracts. Deacon places the
origin of language around two million years ago, whereas most other
authors consider the evolution of language far more recent. The fit
between language and brain, Deacon argues, is the result of the cultural
evolution of invented linguistic forms, not the result of innate
linguistic structures. Derek Bickerton, like Deacon, emphasizes the
functions of language beyond communication. He argues syntax
distinguishes the consciousness of modern humans from that of human
ancestors and of other animals. Language evolved as an adaptation to
function as a representational system, allowing humans to form patterns
of information without waiting for experience. However, all these
matters are controversial.
The study of animal psychology, particularly that of the great apes, is
another subject we will investigate. Sue Taylor Parker compares the
developmental psychology of apes and humans. Although it is certainly
not the case that most new characters are added by terminal addition,
this may occur often enough, and the relation between evolutionary
change and changes in the timing of developmental events may be
important enough, to make comparative developmental studies of human and
ape behavior and mentalities especially
revealing about their evolutionary history.
Chapter VI: ENTITIES OF EVOLUTION - SPECIES AND BEYOND
For centuries biologists and philosophers have been debating the nature
and possible role of various units that may or may not somehow
participate in the evolutionary process. If species, for example, are
mere abstractions, it would seem to make no sense whatever to say that
they can originate, or otherwise evolve. So the modern view, after much
controversy, and with continued resistance, is that they are not
abstractions, but concrete particular things. Supra-individual wholes or
"individuals" in an ontological sense (literature anthologized in
Ereshefsky, 1992). This idea has a misleading similarity to traditional
ideas about societies, ecological communities, and indeed the world
itself being "super-organisms." But it poses a serious alternative to
those who would "reduce" causality to the level of the parts. Thus a
"hierarchical" view of the world (e.g., Salthe, 1985) is opposed to
"reductionist" views (e.g., Williams, 1992). There seems to have been an
over-reaction to group selection with its adapted species. Perhaps there
has been an over- reaction in efforts to revive it (Wilson and Sober,
1989) and even to revive the super-organism notion (Wilson and Sober,
1989). But many interesting possibilities such as species selection have
emerged from considering what goes on at and above the species level
(Eldredge, 1985; Vrba, 1989). The units of selection controversy has led
to much discussion of the role of the various entities that are ranked
at various hierarchical levels (Lewontin, 1970; Hull, 1980). Some of the
more zealous advocates of
sociobiology have given the impression that the only thing that really
matters is the gene. Calling such entities "replicators" gives the
misleading impression that they are things that replicate, rather than
things that are replicated. Be this as it may, biological theory as
generally accepted does attribute different roles, and ones that are
important, to the occupants of different levels. Genes mutate,
chromosomes recombine, organisms reproduce, populations have gene flow,
species speciate. What else such things do, and their relative
importance are still being debated. Also highly controversial is what
may be going on at yet higher levels.=20
This Chapter will reconsider some of these issues, especially the role
of entities at the organismal and
higher levels.=20
Chapter VII: RATES AND KINDS OF EVOLUTIONARY CHANGE - SLOW ACCUMULATION,
SUDDEN LEAPS, OR NEUTRAL DRIFTS
In many ways this Chapter is both an extension of the previous Chapter
and a precursor to the succeeding Chapter. Questions about the rates and
modes of evolutionary change have a long history. It is because of the
observed discontinuities in rates of change, and the recognition that
causal processes operate at a multitude of levels, that many biologists
have begun to see the biological world as intrinsically hierarchical, or
at least best understood with hierarchical models.
Some subjects this Chapter may are punctuated equilibria and coordinated
stasis, evolutionary innovations, challenges to micromutationism, and
chance and the Neutral Theory. investigate, These investigations have
called into question some received views of evolutionary change.=20
Punctuated Equilibrium and Coordinated Stasis
Punctuated equilibrium was the name given by Eldredge and Gould in 1972
[1] to interpretations of an observation: many fossil lineages exhibit
stasis. The origins of morphologically distinct paleontological lineages
are usually geologically instantaneous, and these lineages exhibit
subsequent stability, often for millions of years. The interpretations
include that these 'morphospecies,' defined by morphology, are real
species; that gross morphological change and speciation are often
coincident; and that long-term stasis is a phenomenon requiring
explanation, whether by stabilizing selection or genetic or
developmental constraints. Long-term stasis makes the possibility of
sorting and selection among species relatively more likely, since
populations might speciate or go extinct rather than change in
morphology in response to environmental change. Long term trends in the
history of life might be the result of chance, due to species sorting,
or to species-level properties, such as the ability to speciate to avoid
extinction. Long-term trends might then be decoupled from evolutionary
processes within populations. In particular a disputed question is
whether progressive adaptive trends are common.
A recent topic of investigation is the apparent phenomenon of
coordinated stasis [2]. Groups of coexisting species lineages display
concurrent stability over extended intervals separated by episodes of
abrupt change. Few species become extinct and few immigrants become
established within the region and the period of stability, and species
associations tend to persist. But during the relatively brief intervals
of change, most of the species become extinct, at least locally;
speciation occurs; and immigrants become established in the environment.
Whether coordinated stasis is a generally occurring phenomenon is being
investigated. Some questions include whether and how environmental
tracking and other forms of stabilizing selection occur; the nature of
developmental constraints; how stasis maintains the structuring of
ecological communities; if changes occur gradually, whether episodic
environmental change drives faunal turnover and therefore, whether the
response of fauna to such change may be said to display non-linear
evolution.
The theory of punctuated equilibrium provides just one example of how
paleontology has become better integrated into evolutionary biology as a
whole.
Per Bak's discovery of the principle of Self-Organizing Criticality may
provide the basis of a general theory of complex systems, and Bak has
suggested that the phenomena of punctuated equilibria could be explained
as arising from the intrinsic properties of ecosystems[3]. There is,
however, as yet little consensus on this new theory of evolutionary
change.
Evolutionary Innovations
What are the conditions necessary for major evolutionary innovations?
One of the most remarkable examples of rapid evolutionary change is the
'Cambrian explosion' though there remains controversy over how rapid the
Cambrian explosion actually was, and whether the apparent sudden
appearance of metazoan animals is an artifact of the incomplete fossil
record. In any case several theories have been proposed.
McMenamin and McMenamin [4] propose that ecological chaos resulted from
the origin of predation and the subsequent origin of defenses to
predation.
Jacobs [5] proposes that since selector genes differentiate the segments
of many developing arthropods, this was the ancestral condition of
coelomates. Once these genes may have been closely linked on the genome,
and simple rearrangements may have led to big morphological effects.
Stuart Kauffman [6] suggests that on an adaptive landscape with many
local maxima (a rugged fitness landscape), given a species with low
fitness, big mutations are as likely to be beneficial as small ones;
hence, large morphological changes are possible. Later as the species
reaches higher peaks, big mutations tend to be harmful, and big changes
are unlikely. Also, the landscape may get smoother.
Erwin, Valentine, and Sepkopski [7] propose the 'empty barrel theory':
in the beginning, ways of life are vacant. In the absence of
competition, selection is weak, change is rapid, and there is room for
experiment until the Permian, when the barrel almost fills up.
Challenges to Micromutationism
A common assumption has been that the ultimate sources of variation are
mutations with tiny effects, largely point mutations, and that selection
always controls the direction of evolutionary change; variation never
biases the direction of change. Both these ideas have been challenged.
Kimura argued that mutations with tiny effects, and hence small fitness
benefits, are likely lost to drift; instead, mutations with moderately
large effects are much more likely to be fixed by selection. A review of
the genetics literature by Coyne and Orr [8] showed how surprisingly
little evidence there is of adaptation through micromutation.
In addition, many recent discoveries in molecular genetics pose a
challenge to standard neo-Darwinian models, suggesting the possibility
both of discontinuous change, and the likelihood that variation can bias
the direction of evolutionary change [9]. Among the discoveries are: the
recognition of the importance of 'selfish' DNA; Dover's discovery of
molecular drive in 1982; and the recognition of the overwhelming
importance of lateral gene transfer in the evolution of bacteria.
Additional levels of hierarchy extend below that of the organism; parts
of the genome are subject to sorting and selection and other processes.
Chance and the Neutral Theory
This Chapter may address the role of chance in evolution, as well as the
use of stochastic approaches in evolutionary biology. Genetic drift --
indiscriminate sampling as a result either of sampling error or
selective neutrality -- is a stochastic process. The ideas of drift and
non-adaptive characteristics can be traced back to Delboeuf [10] and
Gulick [11] in the 1870's, and drift played a considerable role in the
work of Sewall Wright [12], who emphasized in particular the role of
drift in maintaining genetic diversity between subpopulations, which
provides in turn abundant variation for natural selection to act on.
However, the discovery of the great importance of genetic drift and
neutral mutation as evolutionary processes came in the late 1960's when
Kimura [13] and King and Jukes [14] separately put forward the idea that
most evolutionary changes at the molecular level are caused by random
genetic drift of selectively neutral or nearly neutral mutations rather
than natural selection. Early studies of protein producing loci, for
example by Lewontin and Hubby [15], had shown levels of variation that
seemed far too high to be maintained by natural selection. Moreover, the
apparent uniformity of molecular evolutionary rates, in comparison with
phenotypic evolution, suggested most substitutions were selectively
neutral or even slightly deleterious.
The Neutral Theory, like Punctuated Equilibrium theory, was and to some
extent still is a matter of controversy. Models of 'constructive neutral
evolution' are becoming increasingly important. These involve biases in
variation, interaction among gene products, and excess capacities: for
example, Ohta's NK-models of protein evolution [16], or Osawa and Jukes'
'codon capture' model for the transition from one genetic code to
another [17].
Chapter VIII: ENVIRONMENTAL TRANSACTIONS - TRANSDUCERS AND TRANSPONDERS
IN BIOCHEMISTRY
Throughout the symposium we have followed several universal themes,
among which are the
following:=20
Development, at once defines and transcends (because of its inherent
teleology) Evolution. Competition, capitalizes on naturally-occurring
Variation (of both the organism and its environment) and produces by
Selection a form which been proven Adaptive, and which therefore
Survives (prevails) into yet another generation - the alternatives being
diminished reproductive success, up to and including genetic cessation,
and Death. Little was made of Death by Darwin (more so by Wallace), but
Death is of course always and everywhere and relentlessly, the agent of
Selection - but sanitized by context for use in Victorian parlors -
"Natural Selection." And Survival, a term much bruited since Darwin
first introduced (via Spencer) "the survival of the fittest." Whether
this term is or is not oxymoronic to the point of having null meaning is
still undecided: Sir Karl Raimund Popper said Yes, and then he aid No,
and then he died. But what is it that survives? This question will have
been addressed in Chapter VI: Entities of Evolution. It is the purpose
of the present Chapter to explore this question further in take the
discussion of Entities into new territory, that is to say the
encompassing Surround, Umwelt, Environment.
So far, we have considered evolution at the level of individuals, of kin
and of species. It is time now to consider modes of evolution which
allow species to cooperate in seeking tandem, reciprocal, adaptations
that enable the survival of one or more species as a group. This form of
evolution is already well known: for example, the wings and beaks of
hummingbirds and the shapes of the flowers that give nectar in return
for pollination. It is called co-evolution, and the mode is called
mutualism. The relationship is straight across, horizontal, with no
particular advantage gained or lost, but the benefit shared equally. Let
it be noted that this process, abundantly present in Nature, cannot
easily be explained by Darwinian Natural Selection in sensu strictu. The
entity is now corporate, rather than individual, and
extra-sociobiological. There is no evidence at the Group level of
altruism by either partner in service solely to the other, or to the
Group as such, either. It is possible, however, to speak of community
ecology in reference to the transaction as a whole, involving mutual
adaptation to features of the shared environment.
Community ecology has been shown to have further, more elaborate,
dimensions such as evolutionary mosaics, in which large congeries of
species co-evolve over large areas and diverse, rather than local,
populations. Further, the mode of co-evolution may vary from so-called
"gene-for-gene" co-evolution, in which one species develops a genetic
change to counter or facilitate the action of a gene in another species
within the congeries, or even a local mutation to meet a local change in
another local species. This can be seen metaphorically as a magnificent
ballet of action and reaction, compensatory, facilitative or defensive,
forever changing and developing new scenarios for life, en masse within
the ecological Surround. There can even occur local duets, or pas de
deux , of evolutionary convergence, or pas de trois, quattre, etc., as
over against the proto-Darwinian, and now perhaps quite rare, pas de
seul of evolutionary change.
It should be noted in passing that there are other forms of community
ecology which are of a negative sort, involving the co-evolution of
mutually beneficial avoidance, rather than cooperation, and even of
mutually beneficial aggression of one "partner" against the other, or
episodic versions of one or another mode, again based upon environmental
necessity. To round out the paradigm, there is also commensalism, in
which one partner gains, but with no cost to the other, and its evil
twin amensalism, in which one partner is destroyed leaving the other
unaffected.
What is newly addressed in this Chapter is the role of the environment
in the evolutionary process. It is interesting to note, as will be
discussed below, the fact that Environment has been a residual category
throughout most post-Darwinian discussions of evolution. The focus has
been on competition and survival, which is dependent upon adaptation,
but adaptation to what has been largely ignored up until the present.
Clearly, it is to the "conditions of existence," put abstractly and with
a kind of dispassionate remoteness; but in fact, adaptation must be
constrained by the very real, very present, very precise requirements
set forth by the natural world, both without ourselves and within. And
that world is forever in Heraklitean flux.
In this perspective, natural selection may be figured as a lottery
turning up variations - not a new idea - but a double lottery, with the
environment turning up variations of its own. Survival comes when the
numbers from both lotteries line up. In popular thought, or rather in
popular unthought, the environment has at least up to now been
considered, seriatim, an annoyance (Dr. Johnson defined a mountain as
"an interruption between two towns"), a religious experience (the
"Sublime" during the Romantic period), recently as a picture-postcard or
a playground, and more lately as a toilet and a toxic dump. In current
apprehension, the
environment is perceived as a source of discord and dismay: global
warming, climate change, hurricanes of unprecedented fury, unparalleled
periods of heat and drought, el Nino and la Nina, and new features
coming soon to a planet near you.
It now seems inescapable that the Environment must be apprehended as not
just like an organism, but as an organism, a life-unit at the highest
level. And as such an entity, it is capable of adaptive reaction to
stimuli produced at constituent lower levels. There is here at least a
metaphoric parallel to the process by how we, as whole organisms at our
level, respond to changes within the constituent systems of our bodies
in an effort to preserve or to restore healthy equilibrium. This
redirection of attention has brought about much interest and
controversy. It is now possible to speak of co-evolution as "evolution
of the biosphere," the Earth itself evolving. This raises in new form
not only the question of evolution, but of the nature of life itself.
For example, the biologist Lynn Margulis contributes this prospective
view of life on our planet:
"What is life?" is a linguistic trap. To answer according to the rules
of grammar, we=20
must supply a noun, a thing. But life on Earth is more like a verb. It
is a material=20
process, surfing over matter like a strange slow wave. It is a
controlled artistic chaos,=20
a set of chemical reactions so staggeringly complex that more than 4
billion years ago
it began a sojourn that now, in human form, composes love letters and
uses silicon
computers to calculate the temperature of matter at the birth of the
universe.
"What is life?" is a scientific and philosophical exploration. Along the
way, it explores the opposite question - what is death?; as well as
delves into the origins of life; Earth's status=20
as a superorganism; the biological connection between programmed death
and sex; the=20
symbiotic evolution of the five organic kingdoms (bacteria, protoctists,
animals, fungi, and plants); the solar basis of our global economy; and
the startling suggestion that life, not just=20
human life, is free to act and has played an unexpectedly large part in
its own evolution.
Heeding Margulis' warning that defining life is a linguistic trap,
presumably because the definer is the thing defined, it is important to
note her claim that Earth is a superorganism. It is to her and to James
Lovelock that we owe the concept of Gaia (variant Ge or Gaea), in Greek
mythology goddess of the earth, with whose own son Uranus she engendered
the Titans and the Cyclopes. The term was first suggested to Lovelock by
William Golding, author of the novel The Lord of the Flies, as a poetic
response to the first photograph of the Earth from space: a mottled blue
and white orb in the midst of a sea of darkness, seeming to be a thing
symmetric and whole unto itself, and pulsing with life.
However poetic and attractive is the concept of an Earth alive with
interactive organisms, and regulated by a single ontological guiding
principle, it is as yet no more than a metaphor and perhaps a crowning
hope for a polluted and damaged planet. Nevertheless, much in human
affairs, including much in science itself, is guided by unproven and
perhaps unprovable metaphors: for example, "survival of the fittest,"
"gene-delimited behaviors," and the like. Gaia may be an illusion, but
at least she is a benign illusion, presiding over daughter-illusions
such as harmony, cooperation, mutualism. The governing metaphor, itself
unproven and therefore perhaps also an illusion, is at least an
alternative offered to strife, competition and greed.
Adopting the notion of an unitary order-of-orders, and one that is both
self-consistent and self-regulating, offers intriguing opportunities for
research, providing an heuristic device and perhaps even an algorithm
for rebuilding the living world as a multiply complex system of
interactive features and organisms working, perhaps only in a final
sense, to produce environmental (i.e., Earth itself) harmony and
equilibrium.
One area of hopeful research has already shown results, and that is what
is called "lateral gene transfer," investigated by Margulis among
bacteria in a limited biome. The preliminary principle was first
demonstrated by Joshua Lederberg (later President of Rockefeller
University, and not coincidentally a current member of H-NEXA), who
demonstrated gene transduction from one bacterium to another, for which
he won the Nobel Prize in 1959.
Margulis' further research tends to demonstrate a surprising fact: that
species can exchange genes with one another, each to the advantage of
the other's advantage. If this work is replicated and substantiated,
whole new worlds open up for research, and old opinions are challenged -
for example, what is the unit of selection that is demonstrated here? It
seems obvious that it is not the individual, but equally obviously it is
not the species, qua species. Something is going in these biospheres
that goes exceeds biomic definitions, and begins to appear as a
life-form that is superorganic . The distant sounds one hears may be the
detonation of a Kuhnian paradigm and the howls of its "community of
practitioners."
Another area of research may be what we are here calling transpondence.
There is no such word. What we have is a term from electronics,
transponder, which is defined as "A radio or radar transmitter-receiver
activated for transmission by reception of a predetermined signal." Its
only use in research at present is in marine biology, where it is
employed for deep-sea exploration. However, it is part of the basic
instrumentation in all types of aircraft. It is a simple device that
transmits and receives digital information about identity, location and
distance between the aircraft and a radio control tower or range
station. It in effect conducts conversations: "I am I, and I am
here...who are you, and where are you?"
An interesting experimental situation in biome-community structure could
be devised on this model, based on pheromones, one the one hand, and
endorphins and enkephalins on the other. Pheromones are the chemical
signals released by sender organisms, and endorphins and enkephalins are
hormones and pentapeptides released by the reception of these by
opiate-mediating center in the brains of receiving organisms. This is a
conceptual model only, posited only to demonstrate one of a range of
possibilities of relationship between and among entities in an
ecological community based upon exchange of information, beginning with
the tantalizing new research involving reciprocal gene-flow.=20
But this is over the horizon of evolutionary biology, and it also marks
the conclusion of our symposium, The Entangled Bank of Evolution: Darwin
and Beyond.
- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -
Envoi. A Puzzlement - Life in a semiotic jungle
Reductionism, Symbiosis, Holism, Wholeness
The American Heritage Dictionary of the English Language, Third Edition
reductionism (noun)
philosophy: holism, organicism, structuralism, functionalism,
reductionism, reductivism
* Reductionism3D holism
---------------
symbiosis (noun)
1. Biology. A close, prolonged association between two or more different
organisms of different
species that may, but does not necessarily, benefit each member.
2. A relationship of mutual benefit or dependence.
* Meaning 2 contradicts the contingent clause of Meaning 1
----------------
holism (noun)
whole: holism, holistic approach, global approach, universalization,
generalization, generality
philosophy: holism, organicism, structuralism, functionalism,
reductionism, reductivism
* Holism3D reductionism
---------------
Webster's New International Dictionary, 2d. edition
* over 4 million words: there is no definition for the word "wholeness"
Some Topics for Chapter VIII to Investigate
The Chapter, Environmental Transactions, covers a lot of ground: the
relationships among evolving populations, the changing biotic
environment, and the changing physical environment: that is, the
conditions of what Darwin called the struggle for existence. Some basic
questions are: what is the relationship between the environment and the
organism? what is a niche? what is a community? How are communities
organized? at what level are they organized? how do they change? what is
the role of competition? what are the roles of cooperation and
mutualism? what is the role of the physical environment? how stable are
communities and ecosystems? is there a "balance in nature"? how
important are history, chance, and contingency to understanding
communities, or can they be explained ahistorically? do ecosystems and
communities have properties over and above those of their component
individual organisms? All of these questions are controversial.
These problems can be addressed in a number of ways: through theoretical
models, through the study of present-day ecosystems, and through the
study of the fossil and geological records of past ecosystems. Some
topics we might address include: the history of the planetary ecosystem,
the Gaia hypothesis, the role of symbiosis in evolution; the interaction
of ecological and genealogical hierarchies, the role of the physical
environment and rare catastrophes in speciation and large-scale
evolutionary trends, the phenomena of coordinated stasis and the
turnover pulse hypothesis; the role of climate change in human
evolution; the idea that there is a higher order structure to ecosystems
that gives them long-term stability; niche theory and competition theory
and in general the development of theoretical ecology.
If we make a distinction between source laws and consequence laws, the
source laws of natural selection are largely the domain of ecology. The
Chapter will help to explain what is known about these laws. It will
address the organization in nature and its causes, and it will
investigate the relative importance of competition, predation, mutualism
and cooperation, changes in the abiotic environment, and other processes
in evolutionary change. It will ask the big question of what nature is
like: is life a war of all against all? a system based on mutual
interdependence? or on peaceful coexistence? are biological communities
stable or unstable? do ecological disturbances, including rare
catastrophes, determine the direction of evolutionary change?
Some provocative, controversial subjects include mutualism and
symbiosis, the Gaia hypothesis, lateral gene transfer,
self-organization, and the economy of nature.
Selection is often regarded as synonymous with competition. Competition
is often redefined to include not only conflict and reliance on a common
resources, but processes that include no interactions at all. For
example, differences in fecundity are called reproductive competition.
As Evelyn Fox Keller writes, "Here 'competition' denotes an operation of
comparison between organisms (or species) that requires juxtaposition
not in nature, but only in the biologist's own mind. This extension,
where 'competition' can cover all possible circumstances of relative
viability and reproductivity, brings with it, then, the tendency to
equate competition with natural selection itself.' [1] However, the
interactions among organisms can determine what is relevant to the
organism's needs, and these interactions can complicate the relationship
between self-interest and competition. In any case there may be more to
life than the Malthusian dynamic of competition through resource
limitation. In particular mutually beneficial interactions among
organisms - mutualism and cooperation -- have become active topics of
investigation [2]. There is ambiguity about terminology, doubt about how
widespread many of these phenomena are, and disagreement about how these
interactions evolved. Many similar problems are posed in the debate over
group selection. For some authors these interactions tend to undermine
the usefulness of self-interest as a basic concept in biology. Fath and
Patten [3] write:
Traditional evolutionary theory depicts survival of the fittest as a
difficult existence based on
danger, conflict and strife. But, another view is emerging of a more
synergistic organization in
which ecosystems on the whole provide hospitable conditions for life.
This world is populated by
organisms mutually adapted and beneficial by virtue of their direct and
indirect interactions. Many
examples of mutualism have been explicitly observed ... and we view
mutualism as an implicit
consequence of indirect interactions and ecosystem organization.
Symbiosis, meaning "living together," was the term created by Anton
DeBary in 1879 to denote associations of different species. An
especially important type of symbiosis is pollination. As Boucher notes,
there were comments about the confounding of symbiosis and mutualism as
early as 1893, but in the most common usage a symbiont can be parasitic.
The ambiguity may result from the absence of terms to describe all the
possible types of interactions among organisms, and the complexity of
these interactions, which are hard to categorize. For example, a
long-standing source of ambiguity has been between benefits to the group
and benefits to the individual. As Boucher points out, if squirrels eat
99% of the acorns but leave 1% in the ground to germinate, they have
likely provided a net benefit to oaks, but certainly no benefit to the
seeds they have eaten.
In endosymbiosis organisms can become incorporated within other
organisms. Lynn Margulis has long advocated the serial endosymbiosis
theory, in which cell organelles originated by hereditary symbiosis. Her
initial arguments that chloroplastids and mitochondria originated as
free living cyanbobacteria and respiring bacteria are now generally
accepted, but the origins of cell undulipodia (such as cilia) from
symbiosis of spirochetes with archaebacteria are still being
investigated.
James Lovelock and Lynn Margulis' Gaia theory [4] looks at the Earth as
a self-regulating system, an evolving "living" system composed of the
biota and their material environment, and predicts that the Earth's
climate and surface and ocean chemistry should have self-regulating
properties. The predominant viewpoint in earth sciences had been that
inorganic forces controlled the Earth's climate; there has since arisen
"Earth systems science," which studies the interactions among the Earth
systems: designated the biosphere, the atmosphere, the geosphere, the
hydrosphere, and the (human) androsphere. James Kirchner [5] provides an
useful critique of the Gaia hypothesis by separating out its various
parts. The idea that biota influences aspects of the abiotic world is
well supported. Whether the biota influences the abiotic environment,
and whether the abiotic environment substantially influences the
evolution of the biota is debated. This is the co-evolutionary
hypothesis. Whether the interaction between the biota and their
environment is characterized by stabilizing negative feedback loops is
debatable. This is the homeostasis hypothesis. Whether the atmosphere is
maintained not just by the biosphere, but for the biosphere, is also
debatable. This is the Daisyworld hypothesis. Lovelock's simple model
describes an Earth-like world populated by light, dark, and gray daisies
that regulate the temperature by their reflectivity. The Daisyworld's
sun, like our Earth's Sun, generates progressively more heat over time,
yet simply
because of the daisies' individual ability to affect the temperature,
which affects the overall ability of the others to grow, the overall
temperature of Daisyworld remains almost constant. Likewise the Earth's
biota may maintain the planet's climate so as to sustain the conditions
necessary for life. A stronger form of this hypothesis is that the biota
manipulates the environment so as to create favorable, or optimal,
conditions for itself. These stronger forms of the Gaia hypothesis lack
general support.
The importance of horizontal, or lateral gene, transfer has received
increasing attention. Koonin and Galperin write about bacteria and
archaea [6], "The most striking conclusions derived by comparison of
multiple genomes from phylogenetically distant species are that the
number of universally conserved gene families is very small and that
multiple events of horizontal gene transfer and genome fusion are major
forces in evolution." But lateral gene transfer is not limited to
prokaryotes. Viruses may cause mutations or even move gene sequences
between bacteria and eukaryotes, including animals. For example, Siew
[7] describes a RNA phage that transducts genetic material between the
bacterium, Salmonella enteriditis, and the mitochondrial genomes of
chicken. Kidwell and Lisch [8] write:
A tremendous wealth of data is accumulating on the variety and
distribution of transposable
elements (TEs) in natural populations. There is little doubt that TEs
provide new genetic=20
variation on a scale, and with a degree of sophistication, previously
unimagined. There are=20
many examples of mutations and other types of genetic variation
associated with the activity
of mobile elements. Mutant phenotypes range from subtle changes in
tissue specificity to=20
dramatic alterations in the development and organization of tissues and
organs... The primary parasitic nature of these sequences during their
invasion of host populations is beyond=20
dispute, but we believe that this does not by any means represent the
whole story. A=20
number of features of both TEs and their hosts can be interpreted as
coadaptations to=20
mitigate or abolish the reduction of fitness due to unbridled
transposition. Furthermore,=20
the number of well documented cases in which TE sequences have been
coopted successfully=20
by the host to provide a useful function is small but is growing
rapidly. We suggest that=20
the process by which elements and their hosts coevolve mutually
beneficial strategies=20
may lend itself to the production of genetic variation that would not
otherwise have arisen.
Several biologists have argued that the ecology of nature and the
economy of man are fundamentally the same: entities producing or
metabolizing, interacting in an economic context. For Vermeij [9] a
central fact of economic interactions is inequality - differences in
metabolism, in kind or number of functions and interactions, and this
inequality is the cause of the directionality in history. The more
dominant individuals disproportionately influence the economy, causing
an increase in energy flux among these dominant individuals. They affect
and restrict other individuals in the economy as well; hence, they
impose a direction on the economy of nature as a whole. Vermeij writes
that there is in the history of life and in human economic history "a
consistent, perhaps stepwise, increase in energy flux, observed both
among ecological dominants and globally at the level of whole
economies." Vermeij however does not call this progress, since progress
implies a value judgment, but he does argue that there is directionality
in history, and that ecological or economic history has predictable
aspects.
Recently there has been great interest in the investigation of sources
of order in natural systems: the study of self-organization, how order
can emerge from systems because of interactions among their components,
often regardless of the physical nature of the systems' parts. In
contrast to natural selection, which involves selection among multiple
entities that differ in fitness according to some criteria external to
the system, self-organization need involve only one system, which
through internal processes moves to some point of stability without
requiring interaction from outside the system. Self-organization can
also be described as the move from a larger to a smaller region of state
space under the control of the system itself, where state space is the
number of possible states available to a system. A stable or persistent
smaller region in state
space is called an attractor. There need be only random change to cause
self-organization to arise, allowing the system to move toward an
attractor. Random fluctuations can cause a system to escape one
attractor and approach another.
Kauffman [10] has developed abstract models of networks that in theory
correspond to genetic regulatory networks and has investigated the
properties that should exist in any real system with the attributes of
the abstract system. Boolean Network or NK models describe a collection
of of N logic gates (AND, OR, NOT, etc.) with K inputs to each gates and
the possible logic functions that can be used with them. These can be
assigned randomly to nodes to create a Random Boolean Network.
Canalyzing functions are functions which are able to force part of the
network into a fixed state given a certain input, and these are thought
closely related to self-organization. A general result is that having
two interconnection for each unit, or allele, leads to optimal adaptive
dynamics. NK models can be extended to NKC Networks and NKCS Networks,
which add additional levels corresponding to other organisms (C) and
other species (S). Weber [11] draws interesting parallels with many of
Sewall Wright's population genetics models, in which a certain level of
gene interaction favors adaptation.
Kauffman describes those systems at the transition between ordered and
chaotic structures as at the "edge of chaos." Small changes can push the
system either into chaotic behavior or into fixed behavior. He suggests
that organizations at the edge of chaos when they become the targets of
natural selection are most likely to result in systems able to
coordinate complex tasks and to adapt.
Criticality is the point where system properties suddenly change.
Self-organizing criticality is the ability of a system to approach a
critical point and maintain itself. Per Bak has developed a general
mathematical theory of self-organizing systems and has proposed,
together with Sneppen and others [12], a simple mathematical model of
macroevolutionary change of interacting species. The environment of each
species is affected by every other evolving species. In their model the
ecology as a whole exhibits 'self-organizing' criticality: "periods of
stasis alternate with avalanches of causally connected evolutionary
changes." Hence, the phenomena of punctuated equilibria, extinctions,
and mass extinctions may be explicable in terms of ecosystem dynamics
and require no external cause. These ideas are controversial.
If there is a common thread to these several topics - mutualism,
endosymbiosis, the Gaia hypothesis, lateral gene transfer,
self-organization - it is that organisms act and interact; they change
and are changed by these interactions.
Chapter IX: SOCIOBIOLOGY AND EVOLUTIONARY PSYCHOLOGY - "THE ETHICS OF
THE GENE"
In 1975, Edward Osborne Wilson, Professor of Zoology and Curator in
Entomology at the Museum of Comparative Zoology of Harvard University,
published Sociobiology: the New Synthesis. It is a monumental work in
evolutionary significance as well as size (697 large pages). Building
upon the work of others in the fields of ethology and population
genetics, it provides a fundamentally new way of understanding
evolution. It provides as well new explanations for a host of other
phenomena in biological and other sciences, including the social
sciences, and in human affairs generally. The impact of this book on the
biological redefinition of human nature itself has been incalculable.
Lest there be no mistake about the principal target of Wilson's
sociobiology being the human world, positing explanations of human
behavior based on extrapolations from observations taken among non-human
species (In Wilson's own domain of expertise, it is ants). Thus, it is
also another example of extreme evolutionary reductionism. Wilson
entitles his first section Social Evolution, in which he enters a domain
well populated from Herbert Spencer forward . But now he gives his
undertaking fresh emphasis by titling his first chapter , The Morality
of the Gene. Here Wilson for the first time applies modern biological
theory to human behavior in its normative aspect, which he finds largely
fictive. Wilson is bound to the task of redeeming for biology not merely
behavior (ethology), but moral behavior. The challenge to do this arose,
as Wilson tells it, because of the biological enigma of altruistic,
self-sacrificing (if not in mortality, then in reproductive potential)
behavior of the individual in behalf of the many: a seeming
contradiction to basic Darwinian doctrine of "survival of the fittest."
The answer, Wilson says, lies in redefining the units of selection, in
effect (though he does not explicitly say as much) both downward and
upward - down to the level of the gene, and up to the level of the kin:
that group of organisms sharing a significant number of genes.
"As more complex social behavior is added to the genes' techniques of
replicating themselves, altruism becomes increasingly prevalent and
eventually appears in exaggerated forms. This brings us to the central
theoretical problem of sociobiology: how can altruism, which by
definition reduces personal fitness, possibly evolve by natural
selection?" Thus, with notable audacity, E.O. Wilson moves from the
level of observable behavior into two new and disparate realms: the
genome and the kin - the molecular and the superorganic. The question of
units and levels of evolution will have already been addressed at this
symposium in Chapters VI and VII -- Entities of Evolution and Kinds of
Evolution. But the moral dimension of evolution is here fresh for
consideration.
Morality and ethics, in this new sociobiological view, are subsets of
ethology refocused through the lens of kin selection and kin inclusive
fitness. That which enables the largest number of adaptive genes to
survive into the next generation constitutes a retroactive conferring of
"moral" and "ethical" qualities. If Darwin's "survival of the fittest"
has been construed as a tautology, then here we encounter the
geno-ethical "right-survival of the right-survivors." Wilson does not
dull the blow to conventional standards and beliefs: "..self-knowledge
[moral behavior] is constrained and shaped by the emotional control
centers in the hypothalamus and limbic system of the brain.... They
evolved by natural selection. That simple biological statement must be
pursued to explain ethics and ethical philosophers, if not epistemology
and epistemologists, at all depths."
Wilson continues, beginning his enlargement of the hegemony of
sociobiology into realms heretofore considered extra-scientific, with
this unusual view of the function of philosophy: "Self-existence... is
not the central question of philosophy. The hypothalamic-limbic complex
automatically denies such logical reduction by countering it with
feelings of guilt and altruism [sic]. In this way the philosopher's own
emotional control centers are wiser than his solipsistic consciousness,
'knowing' that in evolutionary time the organism counts for almost
nothing." Once again, we encounter here the hallmark of late twentieth
century worldview: the reductionistic mode of understanding. But here it
is carried close to a theoretical limit. The rhetorical poles in the
discussion of human and other behavior are these: behaviorism, the mode
of apprehending ethological data, and sociological reductionism.
Behavior-ism was the stock-in-trade of
anthropology, sociology and most schools of psychology. It is this molar
approach Wilson refers to when he speaks of "the stifling grip of the
extreme behaviorists, whose view of the mind of man as a virtually
equipotent response machine" which "was neither correct nor heuristic."
One looks back with a sense perhaps of nostalgia to the anthropologist
Loren Eiseley's dictum (Darwin's Century, 1959) that the human brain is
the universe's only organ of indeterminacy.
The contrary, the reductionist approach to behavior, is to "establish
the lowest taxonomic level at which each character shows significant
intertaxon variation." And that level, again is the gene, or more
precisely DNA; or even more precisely, nucleotides; or more precisely
still, hydrogen bonds; or ever more precisely, complementary bases of
adenine and thymine, or cytosine and guanine; or a pyramidine base and a
purine base; or with ultimate precision in the definition of the origins
of behavior, C4H5N3O and C5H5ON5.
Ne plus ultra.
If sociobiology is reductionist ethology, then it itself implodes to
further reductions: from kin selection to gene selection, from gene
selection to combination and recombination of chemical compounds and
elements. And thus all the ultimate constituents of all behavior of all
human beings alive today, or who have ever lived, or ever will live,
could indeed be placed on the head of a pin.
Here is the epigraph to Sociobiology: the New Synthesis. It is a dialog
from a Hindu epic:
Arjuna to Lord Krishna: Although these are my enemies, whose wits are
overthrown by greed, [do you] see not the guilt of destroying a family,
do [you not] see not the treason to friends, yet how, O Troubler of the
Folk, shall we [if we have] clear sight not see the sin of destroying a
family?
Lord Krishna to Arjuna: He who thinks this Self to be a slayer, and he
who thinks this Self to be slain, are both without discernment: the Soul
slays not, neither is it slain.
This is a fragment of the poem Mahabharata, one of whose episodes
describes the Prince Arjuna, who on the eve of battle, laments the
slaughter that is to follow. The Lord Krishna, with the blue features
that render his divinity, descends to reprimand the young man. It is the
human duty to be a warrior, Krishna says, and Arjuna should not have the
effrontery to think like a god. This poetic lecture on the requirement
of resignation to divine will, without understanding, is called the
Bhagavad-Gita.
This epigraph is no mere exotic ornament to a book of newly-defined
biological evolution. Clearly, it is designed to encapsulate a new
synthesis of modes of understanding of human nature and human destiny.
It is frank. It is monumental. But unlike the Gita, the New Synthesis
provides no over-arching promise of an ultimate meaning to the universe.
It is also dismaying to many people. Consider what is now admissible as
genetically-derived, and therefore presumably inescapable, human
behavior:
"... aggressive dominance systems, with males generally dominant over
females"
"... competition between classes"
"... a genetic predisposition to enter certain classes and to play
certain roles"
"... a single gene [may appear] that is responsible for success and an
upward shift in status...[an]
upward-mobile gene"
"... the evolution of indoctrinability.... The group-selection
hypothesis [sociobiology] is sufficient to
account for the evolution of indoctrinability."
"... endemic warfare and genetic usurpation [as] an effective force in
group selection was clearly
recognized by Charles Darwin" [N.B. the passage adduced from The Descent
of Man does not
demonstrate this assertion.].
"... some of the `noblest' traits of mankind, including team play,
altruism, bravery on the field of
battle [can be shown to be] the genetic product of warfare."
A skeptical observer might wonder if cruelty, despotism and greed do not
belong at the table of gene-driven
imperatives.
The question of human ethical behavior must now be addressed, Wilson
says, through biology. Chapter One of Sociobiology: the New Synthesis is
entitled The Morality of the Gene.
In a section marked "Ethics," Wilson is reaching for the conclusion of
this long argument for sociobiology becoming a master science,
encompassing almost the totality of human knowledge: "Scientists and
humanists should consider together the possibility that the time has
come for ethics to be removed temporarily from the hands of the
philosophers and biologicized.... " and he calls for "the genetic
evolution of ethics." ... "In the first chapter of this book I argue
that the ethical philosophers intuit the deontological canons of
morality by consulting the emotive centers of their own
hypothalamic-limbic system...."
One may wonder why the question of ethics should be dealt with in full
under the aegis of evolutionary anatomical endocrinology or even
sociology , rather than Moral Philosophy or any branch of humanistic
studies. The answer to this question, en passant, is that the humanities
and conventional social sciences have been described in Chapter One as
inevitably to become engulfed by sociobiology: "It may not be too much
to say that sociology and the other social sciences, as well as the
humanities, are the last branches of biology waiting to be included in
the Modern Synthesis." And here, Chapter 27, "In this macroscopic view
of the humanities and social sciences shrink to specialized branches of
biology; history, biography, and fiction are the research protocols of
human ethology; and anthropology and sociology together constitute the
sociobiology of a single primate species.
This of course was not the end of the story of sociobiology and its
offspring, evolutionary psychology. A discussion of its avatars follows.
But at this point, one feels impelled to look again at Wilson's strange
fixation on Indian mysticism for validation of his maximally reductive
materialism. At the close of Chapter Five, he has momentarily seemed
unsure of his stance. In a coda entitled "the field of Righteousness,"
he says: The New Synthesis " [a]bove all...predicts ambivalence as a way
of life in social creatures. Like Arjuna faltering on the Field of
Righteouusness, the individual is forced to make imperfect choices based
on irreconcilable loyalties - between the "right" and "duties" of self
and those of [others].... No wonder the human spirit is in constant
turmoil. Arjuna agonized, " Restless is the mind, O Krishna, turbulent,
forceful, and stubborn; I think it is no more easily to be controlled
than is the wind." And Krishna replied, "For one who is uncontrolled, I
agree the Rule is hard to attain; but by the obedient spirits who will
strive for it, it may be won by the proper way."
One wonders whether, in dreams, Wilsonian sociobiologists are ever
visited by the Blue-Faced God,
sternly calling "Arjuna!"
This of course is not the end of the story of sociobiology and its
offspring, evolutionary psychology. A full discussion of its avatars is
just beginning, here at this symposium and elsewhere. But at this point,
one feels impelled to look again at Wilson's fixation on Indian
mysticism for validation of his extreme reductionistic materialism. At
the close of Chapter Five, he himself has momentarily seemed unsure of
his stance. In a coda entitled "the field of Righteousness," he says:
The New Synthesis " [a]bove all...predicts ambivalence as a way of life
in social creatures. Like Arjuna faltering on the Field of
Righteousness, the individual is forced to make imperfect choices based
on irreconcilable loyalties - between the "rights" and "duties" of self
and those of [others].... No wonder the human spirit is in constant
turmoil. Arjuna agonized, " Restless is the mind, O Krishna, turbulent,
forceful, and stubborn; I think it is no more easily to be controlled
than is the wind." And Krishna replied, "For one who is uncontrolled, I
agree the Rule is hard to attain; but by the obedient spirits who will
strive for it, it may be won by the proper way." [2]
One wonders whether, in dreams, Wilsonian sociobiologists are ever be
visited by the Blue-Faced God, sternly calling "Arjuna!"
Some Background Inforation on Sciobiology and Evolutionary Psychology
Wilson's sociobiology was presented as the beginning of a new discipline
that would encompass and explain the social behavior of all animals,
including humans. One of the essential ways in which its creator saw it
is as distinct from other attempts to discover the biological basis of
human behavior -- for example, the work of Konrad Lorenz -- is indicated
in the first few pages, where Wilson describes the "central theoretical
problem of sociobiology...how can altruism, which by definition reduces
personal fitness, possibly evolve by natural selection?" [1] For Wilson
the solution to the problems of altruism and cooperation, and in turn
the solution to the problem of social behavior, had been provided to a
great extent in the work of two individuals, William Hamilton, with his
mathematical models of inclusive fitness or kin selection [2, 3], and
Robert Trivers, with his model of reciprocal altruism - theories that
Trivers quipped "are designed to take the altruism out of altruism." [4]
Darwin had recognized both the role of the group as the unit of
selection and the influence of kinship on behavior. There was in the
1960s a great trend to rid biology of group selectionist ideas and
replace them with kin-based and game-theoretic models. While Hamilton's
work was not unprecedented - Wright, Malecot, Haldane, and in particular
George Williams, earlier had developed similar mathematical models -
Hamilton provided detailed, rigorous, but straightforward mathematical
models for the evolution of altruism toward near kin [2]. An organism
will extend benefits toward kin only in proportion to the degree it is
related to it; hence, Hamilton's rule: c < rb, where c is the cost of a
behavior to the donor, b the benefit to the recipient, and r is
relatedness; as Haldane remarked, "I'd give my life for 2 brothers or 8
cousins." Such compelling concepts as inclusive fitness -- the fitness
of an organism plus the fitness of its kin adjusted for some coefficient
of relatedness -- and kin selection -- selection that increases
inclusive fitness -- came into common usage.
The evolution of "reciprocal altruistic" behavior among non-kin was
investigated by Trivers [4].=20
There are advantages to cooperation, but the optimal strategy for an
egoist seems to be to cheat - to
accept benefits from others, but to decline to give them. In the long
run, reciprocation should be replaced by the evolution of "reciprocal
altruistic" behavior among non-kin. This was investigated by
Trivers [4]. There are advantages to cooperation, but the optimal
strategy for an egoist seems to be to cheat, to accept benefits from
others, but to decline to give them. In the long run, reciprocation
should be replaced by selfishness. Trivers recognized that situations in
nature are like the game-theory problem known as the "prisoner's
dilemma," in which the best outcome is for A is to cheat while B
cooperates, but the worst outcome is for A and B both to cheat. If A and
B both cooperate, that is the best long term solution for both of them,
but there seemed to be no way this strategy could evolve. Later in 1981
Axelrod and Hamilton found a solution: a tit-for-tat strategy, in which
the organism cooperates the first time, then afterwards follows the lead
of its partner, cooperating if its partner cooperates, cheating if its
partner cheats.
The significance of kin selection is controversial. For some it is just
a rule of thumb, equivalent to standard population genetics models, that
indicates how some interactions among kin contribute to fitness. But for
others kin selection was revolutionary. Richard Dawkins borrowed
Hamilton's phrase, "the selfish gene," and William's gene selectionism
to create a vision of evolution in which genes are the units of
selection [5]. For Dawkins, perhaps the most important implication of
kin selection is that genes are the beneficiaries of selection:
evolution is for the genes. For example, kin selection may favor a gene
or genes that cause(s) a sterile worker honeybee to give up reproduction
and possibly to give up her life to defend her hive; it is the genes
that will survive; the organism is disposable -- or so it seems in this
view of life.
For others kin selection is in fact a form of a group selection, despite
the tendency to present it in opposition to group selection. The
original applications of kin selection to explain social behavior --
Hamilton's relatedness asymmetry hypothesis for the origin of
eusociality in bees and ants, in which
central importance was attached to the _ relatedness of females workers
among haplodiploid hymenopterans -- is not generally accepted [6]. Kin
selection models for the evolution of insect sociality do have majority
support, but some biologists dissent - Aoki, Ito, Michener, Lin, and so
on. In light of E.O. Wilson's remarks, "Let me say that sociobiology,
the study of the biological basis of social behavior, was never in
dispute when applied to animals," [7] it is worthwhile noting that there
is controversy even in insect sociobiology.
The publication of Wilson's great work was announced on the front page
of the New York Times magazine. Wilson won the Pulitzer Prize and the
National Medal for Science. The Animal Behavior Society voted
Sociobiology: The New Synthesis the most important book on animal
behavior ever published. The ideas of human sociobiology were spread
through the popular media. Human sociobiology and its descendents have
achieved a small but deeply entrenched place in academia, and a rather
large place in the popular imagination.
However, in addition to the honors and publicity, there was opposition
to sociobiology: petitions and demonstrations against the rebirth of
social Darwinism, and detailed critiques. The criticisms provided by
Stephen J. Gould and Richard Lewontin in Natural History and the New
York Review of Books had particular force, since they had prestige equal
to Edward O. Wilson's.=20
Lewontin's critique of sociobiology [9] is especially incisive, since he
argues that some fundamental errors of sociobiology are errors of the
"adaptationist program" in general. That is, sociobiology's problems are
not just with humans but are more general. The adaptationist program,
according to Lewontin, may include
an attempt to identify the problems an organism is trying to solve, the
identification of optimal solutions, and comparison of the solutions
with optimal practice. Or the program can start with some part of the
phenotype of the organism, assume it is a solution to a problem, and
then identify the problem it solves. Lewontin suggests that some basic
problems with adaptationist theory include fundamental errors of
description, confusion about heritability studies, and adaptive
story-telling. Problems with description include arbitrary agglomeration
(what should be studied), reification (whether what we give a name is a
real thing affected by selection), anthropomorphism and conflation
(whether behaviors in humans and animals that we assign the same name
are in fact linked), and reductionism and confusion of levels (whether
social behavior is the sum of individual behavior). In general
sociobiologists confuse ideological constructions like slavery and
tribalism with material objects. This is an error of reification, since
selection acts on material objects, not mental constructs.
Sociobiologists conflate unrelated phenomena, with different functions
and causes, named by the same word - conflating political aggression,
which has political and economic causes, with instinctual aggression in
animals, or, as Barash did, conflating, coercive sexual reproduction in
ducks with rape in humans. This rhetorical legerdemain might be called
semantic phyletic reductionism and phyletic semantic constructionism.
Adaptive story-telling, Lewontin argues, is flawed, by "progressive
ad-hoc optimization." The premise that a trait is an optimal solution to
some problem is protected from falsification because a trait is examined
essentially in isolation, and if one adaptive story fails, another takes
its place without the overall premise of optimality being questioned.
Adaptive story-telling is also flawed by "imaginative reconstruction."
(See our suggested usage immediately above.) Sociobiologists create
scenarios about why traits should be favored by natural
selection, but they do so without experimental proof. Lewontin also
discusses several scenarios in
which non-optimality is expected, because of random drift, selection on
correlated characters, developmental noise, and other processes.
- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -
- - - - - - - - - - - - - - - - - - - - - - - - - - - - -
In the years since the publication of Wilson's Sociobiology, a new
discipline has arisen called evolutionary psychology. It is hardly
unprecedented. Darwin's notebooks and his published work, especially The
Expression of the Emotions in Man and Animals and The Descent of Man,
reveal his thoughts on the ontogeny and phylogeny of behavior, emotional
expression, sexual selection, the evolution of morals, and the
inheritance of behavior (10). Evolutionary psychology tries to step back
from sociobiology's attempt to explain social behavior, and instead
first tries to explain human "mentalities." Pinker summarizes the "key
idea" of evolutionary psychology: The mind is a system of organs of
computation, designed by natural selection to solve the kinds of
problems our ancestors faced in their foraging way of life, in
particular, understanding and outmaneuvering objects, animals, plants,
and other people... The mind is organized into modules or mental organs,
each with a specialized design that makes it an expert in one arena of
interaction with the world. The modules' basic logic is specified by our
genetic program. Their operation was shaped by natural selection to
solve the problems of the hunting and gathering life led by our
ancestors in most of our evolutionary history. [11]
A central method for evolutionary psychology is "reverse engineering":
cognitive psychology describes an information processing mechanism, and
evolutionary psychologists describe what it was an adaptation for in our
hunter-gatherer ancestors. "There is no reason that reverse-engineering
guided by evolutionary theory should not bring about insight about the
rest of the mind." [12] Here is a classic example of evolutionary
psychology "reverse engineering." Four cards are set on a table with a
letter on one side and a number on another, and human subjects are shown
one side of each card:
card 1: D card 3: 3
card 2: F card 4: 7
They are then asked which card they need to turn over to falsify or
verify the hypothesis that "If a card has a D on one side, it has a 3 on
the other." The correct answer is to turn over 1 and 4, since what
matters is that a "D" card must have a 3 on the other side, and the "7"
card can't have a D on the other side. However, most people answer 1, or
1 and 3. But the problem is restated, "You are a bartender, and you need
to enforce the rule, `If a person is drinking beer, he must be eighteen
or older,'" and is given cards,
card 1: beer card 3: age 25
card 2: Coke card 4: age 16
in which the person's beverage order is one side and the person's age is
on the other. If the subjects are asked, "Which cards do you have to
turn over to make sure the drinking-age rule is being enforced?" most
perform the task correctly. Since other experiments showed it was not
just a matter of real world objects replacing abstract objects, Cosmides
suggests this phenomenon is evidence of a brain module specialized for
detecting cheaters. Since earlier work in sociobiology, notably by
Trivers, has argued for the importance of reciprocal altruism in the
evolution of social relationships and in particular the evolution of
tit-for-tat strategies, Cosmides thinks it highly plausible that a
mental organ for detecting cheating in conspecifics should evolve [13].
One study often cited as among the best cases for evolutionary
psychology is Daly and Wilson's study of violence against step-children.
Evolutionary psychology predicts that men have no interest in the
survival of genetically unrelated step-children. It is against their
genetic self-interest. Therefore, Daly and Wilson predict the males
should be much more likely to harm step-children than their own
children. As predicted, abuse is seven times higher; fatal abuse is 100
times higher. [14]
- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -
- - - - - - - - - - - - - - - - - - - - - - - - - - - - - -
Group selection, banished by Hamilton, Maynard-Smith, George Williams
and others during the 1960's, has slowly been returning to evolutionary
biology. It may help to understand the evolution of human sociality,
since presumably so much of human evolution occurred in small groups.
Sober and Wilson in Unto Others: The Evolution and Psychology of
Unselfish Behavior [15] investigate both evolutionary and psychological
altruism. They clearly distinguish evolutionary altruism, which involves
mechanisms that affect fitness and is independent of motivation, from
psychological altruism, which involves complicated internal motivations,
but they believe the former can help explain the latter. In particular,
they argue for a return to a consideration of the importance of group
selection, group-level adaptation, and group-level functionalism. This
leads to the possibility that unconditional altruism could evolve. Kin
selection models predict that individuals surrender their self-interest
only in proportion to their genetic relatedness to the beneficiary.
Group selection models predict that a group of unrelated individuals can
behave as an adaptive unit, even without close kinship. As a result,
Sober and Wilson consider motivational pluralism likely: both true
egoism and altruism exist. They write:=20
As the most facultative species on earth, human beings appear willing
and able to=20
span the full spectrum from mercilessly exploiting their social partners
to sacrificing=20
their lives for others. This stunning plasticity of behavior must
somehow be orchestrated=20
by proximate mechanisms that probably include altruistic ulterior
motives, but certainly
include more, such as emotions, moral principles, and other mechanisms
that we deliberately
set to one side to make headway on the subject of psychological
altruism. To the extent that behavior has evolved by group selection,
multilevel selection theory will be required to=20
understand the full architecture of human motivation.
Boyd and Richerson have developed models of cultural evolution. Natural
selection on cultural variation can produce adaptation. Sober and Wilson
have developed these ideas to show how powerful social norms can be
established through adaptive processes, when systems of rewards and
punishments are established. Different societies will evolve to
different internally stable configurations. Neutral variation and
cultural random drift can occur, as can cultural group selection. (see
Chapter VII: Rates and Kinds of Evolutionary Change). Theses processes
can produce group-level functional organization, but Sober and Wilson
also point that they can promote nonfunctional and dysfunctional
behaviors that reduce the welfare of individuals and of society as a
whole.
----------------------------------------------------------------------
REFERENCES
Introduction
1. Cool Memories, ch. 4 (1987; tr. 1990)
2. In Critique, no. 419 (Paris, April 1982; repr. in The Postmodern
Condition: A Report on Knowledge, 1979; rev. 1986)
3. Waiting for Godot, 1954, Grove Press
Chapter I: ORIGINS OF LIFE - ON EARTH AND IN THE COSMOS
Cairns-Smith, A.G. Seven Clues to the Origin of Life. 1995. Discusses
the "clay life" hypothesis, in which the first life had a mineral
substrate.
De Duve, Christian. Vital Dust. Address the origins of life through the
origin of consciousness. Argues that life is chemically inevitable given
the proper initial conditions, such as existed in the prebiotic Earth.
Eigen, Manfred. Steps Towards Life : A Perspective on Evolution.
Discusses the possible role of self-organizing hypercycles -
autocatalytic chemical networks - in the origins of life.
Jakovsky, Bruce. The Search for Life on Other Planets. A review of the
search for life oustide the Earth.
Kauffman, Stuart. The Origins of Order : Self-Organization and Selection
in Evolution. The most prominent work in the application of complex
systems theory to the origins of life and evolutionary
biology. Kauffman argues that self- organization plays an essential role
in the origin of complex systems, including the origins of life and its
subsequent evolution: complex systems, counter-intuitively, can suddenly
exhibit order, and this tendency toward "order for free" contributes to
understanding the origins of life.
Schopf, J. William. Cradle of Life : The Discovery of Earth's Earliest
Fossils. Discusses the author's discovery of the oldest fossils in the
Apex Chert of Western Australia, the conditions of the early Earth, and
the investigation into origin of the earliest cells. In addition, Schopf
discusses the possible traces of life discovered in Martian rocks.
Shapiro, Mark. Planetary Dreams : The Quest to Discover Life Beyond
Earth. A review of the search for life beyond the Earth; addresses, in
particular, the biochemical conditions elsewhere in our solar system.
Taylor, Michael Ray. Dark Life : Martian Nanobacteria, Rock-Eating Cave
Bugs, and Other Extreme
Organisms of Inner Earth and Outer Space. Discusses "extremophiles" -
microbes that live in extreme environments like ice or hot water or deep
in the earth, and the Martian meteorite that may contain fossil traces
of Martian nanobacteria.
Chapter II: EVOLUTION - THEORY AND CONCEPT, PAST AND PRESENT
Bender,Bert (1996): The Descent of Love: Darwin and the Theory of Sexual
Selection in American
Fiction, 1871-1926. University of Pennsylvania Press, Philadelphia. xvi
+ 440 pages.
Bowler,Peter J (1984): Evolution: the History of an Idea. University of
California Press,
Berkeley. xiv + 412 pages.
Gayon,Jean (1998): Darwinism's Struggle for Survival: Heredity and the
Hypothesis of Natural
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Greene,John C (1981): Science, Ideology, and World View: Essays in the
History of Evolutionary
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Landau,Misia (1991): Narratives of Human Evolution. Yale University
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202 pages.
Smocovitis,Vassiliki Betty (1992): Unifying biology: the evolutionary
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Chapter III: "EVOLUTION" AND "PROGRESS" - THE HISTORY OF SOCIAL IDEAS
1. Variations In Value Orientations, Kluckhon, Florence and Strodtbeck,
Fred L. (1961)
2. David Ricardo, Principles of Political Economy and Taxation (1817)
3. On Liberty (1859)
4. Le Comte de Saint-Simon (Claude Henri de Rouvroy), The New
Christianity (1825)
5. A New View of Society (1788)
6. God and the State (published posthumously in 1871; reprinted in
Bakunin on Anarchism, edited
by Sam Dolgoff (1980)
7. The Communist Manifesto (with Friedrich Engels),1848); Das Kapital
(1867)
8. The Subjection of Women, ch. 4 (1869)
9. An Essay on the Principle of Population (1798)
10. Melville wrote Billy Budd, Foretopman in 1891, but it was published
posthumously only in 1924. The intended sequence of chapters is in
question.
Chapter IV: EVOLUTION BEYOND BIOLOGY
Ghiselin, Michael T (1992): Biology, economics, and bioeconomics. Chap.
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Radnitzky,G) Paragon
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Campbell,DT (1960): Blind variation and selective retention in creative
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(Ed: Schilpp,PA) Open Court, La Salle, 413-463. [History in extenso.].
Ghiselin, Michael T. (1994): The Origin of Vertebrates and the Principle
of Succession of Functions.
Genealogical Sketches by Anton Dohrn. 1875. An English translation from
the German,
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Hirshleifer,Jack (1985): The expanding domain of economics. Amer. Econ.
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Mayr, Ernst (1960): The emergence of evolutionary novelties. In: The
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Mokyr,Joel (1990): The Lever of Riches: Technological Creativity and
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Witt,Ulrich (1999): Evolutionary economics and evolutionary biology.
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Chapter V: EVOLUTION AMONG THE PRIMATES - OUR HOMINID ORIGINS AND OUR
ANCESTORS
Bickerton, Derek. Language and Species. 1992. Speculates on the
evolutionary events that led to the evolution of language, the primary
function of which, Bickerton argues, is representing the world, rather
than communication.
Deacon, Terrence. The Symbolic Species. 1997. Argues that language arose
from symbolic thinking when humans became hunters and needed to develop
group cooperation.
Gottlieb, Gilbert. Synthesizing Nature-Nurture: Prenatal Roots of
Instinctive Behavior. 1997. Based
on field observations and experiments on ducks, Gottlieb suggests how
behavior and psychology can mediate the evolution of species, including
humans.
Oyama, Susan. The Ontogeny of Information. 1985. An important work in
the developmental systems approach to evolution, which attempts to get
beyond nature-nurture controversies by arguing that is developmental
systems that evolve. Organized developmental systems that interact with
their environments, rather than genes, are what reproduce and evolve.
Parker, Sue Taylor, and McKinney, Michael. Origins of Intelligence : The
Evolution of Cognitive Development in Monkeys, Apes, and Humans.
Comparative psychology of humans and their relatives. The authors
emphasize the role of development and changes in the timing of
development (heterochrony) in evolution.
Schwartz, Jeffrey. Sudden Origins : Fossils, Genes, and the Emergence of
Species. A review of human evolution, pointing out the evidence for
punctuated equilibrium in hominid lineages and the role of changes in
regulatory genes in evolution..
Stringer, Christopher, and McKie, Robin. African Exodus : The Origins of
Modern Humanity. An account of the "Out of Africa" model of the origin
of modern humans, in which Homo sapiens originated relatively recently
in Africa and spread throughout the world less than 100,000 years ago,
replacing all other human populations. The dispute between the "Out of
Africa" model and the less popular "Multi-regional hypothesis" carries
some social significance. Among those biologists who recognize
subspecies as useful categories, they are usually defined as distinct
evolutionary lineages within species. Hence, advocates of the
"multi-regional hypothesis" argue that the commonly identified human
races are biologically meaningful categories.
Tattersall, Ian. Becoming Human: Evolution and Human Uniqueness. Looks
at the evolution of the unique characteristics of the human species,
emphasizing the complexities and uncertainties of the questions of human
evolution.
Chapter VI: ENTITIES OF EVOLUTION - SPECIES AND BEYOND
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Lloyd,Elisabeth A (1989): A structural approach to defining units of
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Vrba,Elisabeth S; Eldredge,Niles (1984): Individuals, hierarchies, and
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146-171.
Williams,George C (1992): Natural Selection: Domains, Levels, and
Challenges. Oxford University Press, New York. x + 208 pages.
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Chapter VII: RATES OF EVOLUTIONARY CHANGE - DUSTY EONS OR SUDDEN JUMPS
[1] Eldredge, Niles, and Stephen J. Gould. 'Punctuated equilibria: An
alternative to phyletic gradualism.' In Models in Paleobiology, ed. by
T.J.M. Schopf, pp. 82-115.
[2] Brett, C.E., L.C. Ivany, and K.M. Schopf. 'Coordinated stasis: An
overview.' Palaeogeography, Palaeoclimatology, Palaeoecology 127 (Dec.
20, 1996): 1-20.
[3] Bak, Per. 1996. How Nature Works: The Science of Self-Organized
Criticality.
[4] McMenamin, M. A. S., and D. L. S. McMenamin. 1990. The Emergence of
Animals: the Cambrian Breakthrough.
[5] Jacobs, D.K. 'Selector genes and the Cambrian radiation of
Bilateria.' Proceedings of the National Academy of Science 87:4406-4410.
[6] Kauffman, S.A. 'Cambrian explosion and Permian quiescence:
implications of rugged fitness landscapes.' Evolutionary Ecology
3(1989):274-281.
[7] Erwin, D.H., J.W. Valentine, and J.J. Sepkopski, Jr. 'A comparative
study of diversification events: the early Paleozoic versus the
Mesozoic.' Evolution 41 (1986):1177-1186.
[8] Orr, H. A. and J. A. Coyne. The genetics of adaptation revisited.
American Naturalist. 140(1992): 725-742.
[9] Campbell, K.S.W. and M.F. Day, ed. 1987. Rates of evolution.
[10] Delboeuf, J.R.L. 'Le math=82matique et le transformisme. Une loi
math=82matique applicable85 la th=82orie du transformisme. ' La Revue
Scientifique de la France et de l'=90tranger series 2, v. 12 (1877):
669-680. Also published in Kosmos 2 (1877): 105-127. The well-known
psychologist, Delboeuf, shows how disadvantageous transformations can
occur through chance processes.
[11] Gulick, J.T. 'On diversity of evolution under one set of external
conditions.' Journal of the Linnaean Society of Zoology 11(1872):
496-505. Gulick writes, suggesting that variation within species and
speciation itself may be non-adaptive: 'If we suppose separation without
a difference of external circumstances is a condition sufficient to
ensure variation, it renders intelligible the fact that in nearly allied
forms on the same island, the degree of divergence in type is in
proportion to the distance in space by which they are separated. The
difference between two miles and ten miles makes no change in climate;
but it is easy to believe that it is the measure of a corresponding
difference in the time of separation. In forms that differ more
essentially, the separation may have been as complete and as
long-continued in the case of those, which now inhabit one valley as in
the case of those, which are separated by the length of an island. When
a wide degree of divergence has been established, hybridation would be
precluded. '
[12] Provine, W.B. 1986. Sewall Wright and Evolutionary Biology.
[13] Kimura, M. 1983. The Neutral Theory of Molecular Evolution.
[14] King, J.L., and T.H. Jukes. 'Non-Darwinian evolution.' Science 164
(1969): 788-798.
[15] Lewontin, R.C., and J.L. Hubby. 'A molecular approach to the study
of genic heterozygosity in natural populations.' Genetics 54 (1966):
595-600.
[16] Ohta, T. 'The meaning of near neutrality at coding and non-coding
regions.' Gene 205:261-267.
[17] Osawa, S., T.H. Jukes, K. Watanabe, and A. Muto. 'Recent evidence
for evolution of the genetic code.' Microbiological Review 56:229-264. ,
Per. 1996. How Nature Works: The Science of Self-Organized Criticality.
Brett, C.E., L.C. Ivany, and K.M. Schopf. 'Coordinated stasis: An
overview.' Palaeogeography, Palaeoclimatology, Palaeoecology 127 (Dec.
20, 1996): 1-20.
Campbell, K.S.W. and M.F. Day, ed. 1987. Rates of evolution.
Delboeuf, J.R.L. 'Le mathematique et le transformisme. Une loi
mathematque applicable a la theorie du transformisme. ' La Revue
Scientifique de la France et de l'Etranger series 2, v. 12 (1877):
669-680. Also published in Kosmos 2 (1877): 105-127.
Eble, G.J. 'On the dual nature of chance in evolutionary biology and
paleobiology.' Paleobiology 25 (Winter, 1999): 75-87. Clarifies the
usage of chance in evolutionary biology. Chance has both a statistical
meaning and a meaning that refers to natural selection: 'The former
implies a combination of indiscriminate sampling and unpredictability
due to multiple causes; the latter codifies independence from adaptation
and the directionality imposed by natural selection.'
Eldredge, Niles. 1985. Time frames: the rethinking of Darwinian
evolution and the theory of punctuated equilibria.
Eldredge, Niles, and Stephen J. Gould. 1972. 'Punctuated equilibria: An
alternative to phyletic gradualism.' In Models in Paleobiology, ed. by
T.J.M. Schopf, 82-115. Gould and Eldredge propose that through most of
their existence species exhibit morphological stasis.
Erwin, D.H., J.W. Valentine, and J.J. Sepkopski, Jr. 'A comparative
study of diversification events: the early Paleozoic versus the
Mesozoic.' Evolution 41 (1986): 1177-1186.
Gulick, J.T. 'On diversity of evolution under one set of external
conditions.' Journal of the Linnaean Society of Zoology 11(1872):
496-505. The beginning of the study of random drift in evolution.
Jacobs, D.K. 'Selector genes and the Cambrian radiation of Bilateria.'
Proceedings of the National Academy of Science 87:4406-4410.
Jablonski, D., and D.J. Bottjer, 1990. 'The origin and diversification
of major groups: Environmental patterns and macroevolutionary lags.' In:
P.D. Taylor and G.P. Larwood, eds. 1990. Major Evolutionary Radiations,
p. 17-57.
Kauffman, S.A. 'Cambrian explosion and Permian quiescence: implications
of rugged fitness landscapes.' Evolutionary Ecology 3(1989): 274-281.
Kimura, M. and T. Ohta. 1971. Theoretical Aspects of Population
Genetics.
Kimura, M. 1983. The Neutral Theory of Molecular Evolution.
King, J.L., and T.H. Jukes. 'Non-Darwinian evolution.' Science 164
(1969): 788-798.
Lewontin, R.C., and J.L. Hubby. 'A molecular approach to the study of
genic heterozygosity in natural populations. II. Amount of variation and
degree of heterozygosity in natural populations of Drosophila
pseudoobscura.' Genetics 54 (1966): 595-609.
McMenamin, M. A. S., and D. L. S. McMenamin. 1990. The Emergence of
Animals: the Cambrian Breakthrough.
Nitecki, M.H., ed. 1990. Evolutionary Innovations.
Nitecki, M.H., and A. Hoffman. 1987. Neutral Models in Biology.
Ohta, T. 'The meaning of near neutrality at coding and non-coding
regions.' Gene 205:261-2 67..Osawa, S., T.H. Jukes, K. Watanabe, and A.
Muto. 'Recent evidence for evolution of the genetic code.'
Microbiological Review 56:229-264. Directional mutational pressure and
drift can cause changes in the genetic code.
Orr, H. A. and J. A. Coyne. The genetics of adaptation revisited.
American Naturalist. 140 (1992): 725-742.
Provine, W.B. 1986. Sewall Wright and Evolutionary Biology.
Schopf, T.J.M., ed. 1972. Models in Paleobiology.
Taylor, P.D. and G.P. Larwood, eds. 1990. Major Evolutionary Radiations.
Chapter VIII: ENVIRONMENTAL TRANSACTIONS - TRANSDUCERS AND TRANSPONDERS
IN BIOCHEMISTRY
[1] Keller, E.F. 'Competition: Current Usages' In: Keywords in
Evolutionary Biology, ed. Lloyd, E., and E.F. Keller.
[2] Boucher, D.H. 'Mutualism and cooperation.' In: Keywords in
Evolutionary Biology, ed. Lloyd,
E., and E.F. Keller.
[3] Fath, B.D. and B.C. Patten 'Network synergism: Emergence of positive
relations in ecological
systems.' Ecological Modelling 107 (1998): 127-143.
[4] Brown, P.R.F. 'The Gaia hypothesis.' 1996.
Web:http://magna.com.au/~prfbrown/gaia_int.html.
[5] Kirchner, J.W. 'The Gaia hypothesis: can it be tested?' Reviews of
Geophysics 27:2 (1989) 223-235.
[6] Koonin, E.V., and Galperin, M.Y. 'Prokaryotic genomes: the emerging
paradigm of genome-based microbiology.' Current Opinion in Genetics and
Development 7 (Dec. 1997): 757-763.
[7] Siew, R.S. "Microbiology: Millennium bug." Nature. 398 (Apr. 1,
1999): 376.
[8] Kidwell, Margaret, and Damon Lisch. Transposable elements as sources
of variation in animals and plants. Proceedings of the National Academy
of Science 94 (July 1997):7704-7711.
(Colloquium Paper)
[9] Vermeij, G.J. 'Inequality and the directionality of history.'
American Naturalist 153 (1999):243-253.
[10] Kauffman, S. The Origins of Order - Self-Organization and Selection
in Evolution. 1993.
[11] Weber, B.H. The origins of order: self-organization and selection
in evolution.' Biology and Philosophy 13 (Jan. 1998): 133-144.
[12] Sneppen K, et al. 'Evolution as a self-organized critical
phenomenon.' Proceedings of the National Academy of Science 92 (May 23,
1995): 5209-5213.
Boucher, Douglas H. Mutualism. 1985. Defines mutualism and argues for
the importance of mutualistic interactions in nature.
Boucher, D.H. 'Mutualism and cooperation.' In: Keywords in Evolutionary
Biology, ed. Lloyd, E., and E.F. Keller.
Brandon, Robert N. Adaptation and Environment. 1990. Analyzes the
relation of the environment and selection, distinguishing among the
external, ecological, and selective environments.
Fath, B.D. and B.C. Patten B.C. 'Network synergism: Emergence of
positive relations in ecological systems.' Ecological Modelling 107
(1998): 127-143.
Keller, E.F. 'Competition: Current Usages' In: Keywords in Evolutionary
Biology, ed. Lloyd, E., and
E.F. Keller.
Kauffman, S. The Origins of Order - Self-Organization and Selection in
Evolution. 1993.
Kidwell, M., and D. Lisch. Transposable elements as sources of variation
in animals and plants. Proceedings of the National Academy of Science 94
(July 1997):7704-7711. (Colloquium Paper)
Kirchner, J.W. 'The Gaia hypothesis: can it be tested?' Reviews of
Geophysics 27:2 (1989) 223-235.
Koonin, E.V., and M.Y. Galperin. 'Prokaryotic genomes: the emerging
paradigm of genome-based microbiology.' Current Opinion in Genetics and
Development 7 (Dec. 1997): 757-763.
Lewontin, R. C. "Organism and environment." Learning, Development, and
Culture, ed. H.C. Plotkin, 1982. Argues that organisms are not passive
objects shaped by their environments but instead actively construct
their environments.
Margulis, Lynn. Symbiotic Planet : A New Look at Evolution. Discusses
the theory of Serial Endosymbiosis and the Gaia hypothesis and their
relationships - how symbiosis is a key toward understanding the
evolution of multicellularity, the evolution of sex, the emergence of
life on land, and homeostatic balance of the Earth's ecosystems.
Raup, David. Extinction : Bad Genes or Bad Luck? 1992. Raup discusses
the role of mass extinctions in evolution, and the role of periodic
meteor impacts in causing them. The present pattern of diversity of
animal life on Earth may be largely a consequence of past mass
extinctions.
Siew, R.S. "Microbiology: Millennium bug." Nature 398 (Apr. 1, 1999):
376.
Sneppen K; Bak, P; Flyvbjerg H.; and Jensen, M.H. 'Evolution as a
self-organized critical phenomenon.' Proceedings of the National Academy
of Sciences 92 (May 23, 1995): 5209-5213.
Vermeij, G.J. Inequality and the directionality of history. American
Naturalist 153 (1999):243-253.
Weber, B.H. The origins of order: self-organization and selection in
evolution.' Biology and Philosophy 13 (Jan. 1998): 133-144.
Chapter IX: SOCIOBIOLOGY AND EVOLUTIONARY PSYCHOLOGY - "THE ETHICS OF
THE GENE"
[1.] Sociobiology: the New Synthesis (1975), infra.
[2.] Ibid., 129
Axelrod, Robert. & Hamilton, William D. `The evolution of cooperation,'
Science 211, 379-403
(1981).
Alexander, Richard D. Darwinism and Human Affairs. 1979. One of the most
important contributions to sociobiology and evolutionary ethics in the
1970's.
Barkow, Jerome; Cosmides, Leda; Tooby John. The Adapted Mind. 1992. An
importantcollection of articles on evolutionary psychology, which argues
that the brain consists of distinct modules specialized for adaptive
tasks.
Daly, M., & Wilson, M. I. (1996). Violence against step-children.
Current Directions in Psychological Science, 5, 77-81.
Dawkins, Richard. (1976) The Selfish Gene.
Hamilton, William. D. (1964) `The genetical evolution of social
behavior, I and II,' Journal of Theoretical Biology, 7, 1-52.
Hamilton, William D. (1972) `Altruism and related phenomena, mainly in
social insects.' Annual Review of Ecology and Systematics 3: 193-232.
Hamilton, William. D. (1995) The Narrow Road to Geneland: The Evolution
of Social Evolution.
Collection of his papers.
Kitcher, Philip. Vaulting Ambition. 1987. A book length critique of "pop
sociobiology," arguing that pop sociobiology has not made its case.
Lewontin, Richard C. (1979) `Sociobiology as an adaptationist program.'
Behavioral Science 24: 5-14.
Nitecki, Michael H., ed. Evolution and Ethics.
Pinker, Steven. (1997) How the Mind Works.
Sober, Elliott, and Wilson, David S. (1998) Unto Others : The Evolution
and Psychology of Unselfish Behavior.
Ruse, Michael and Wilson, Edward O. "Moral philosophy as an applied
science." Philosophy (1986) 61:173-192. Ruse and E.O. Wilson defend the
biological origin of ethical beliefs and sentiments and look forward to
the possibility that biology can someday apply "correctives ... to
formulate more enduring moral codes."
Sober, Elliott and Wilson, David Sloan. Unto Others: The Evolution and
Psychology of Unselfish Behavior, 1998. A study of evolutionary and
psychological altruism and cooperation, which in particular argues that
group selection could be an effective mechanism toward the evolution of
altruistic behavior.
Trivers, Robert. (1971) `The evolution of reciprocal altruism.'
Quarterly Review of Biology 46, 35-57.
Wilson, Edward O. (1975) Sociobiology: the New Synthesis.
Wilson, Edward O. (1995) Naturalist.
CHAPTER IX: Some Background Inforation on Sciobiology and Evolutionary
Psychology
[1.] Sociobiology: the New Synthesis (1975), infra.
[2.] Hamilton, William. D. (1964) `The genetical evolution of social
behavior, I and II,' Journal of Theoretical Biology, 7, 1-52.
[3.] Hamilton, William D. (1972) `Altruism and related phenomena, mainly
in social insects.'
Annual Review of Ecology and Systematics 3: 193-232.
[4.] Trivers, Robert. `The evolution of reciprocal altruism.' Quarterly
Review of Biology 46, 35-57
(1971).
[5.] Dawkins, Richard. (1976) The Selfish Gene.
[6.] Kasuya, E. 1982. Factors governing the evolution of eusociality
through kin selection.
Researches on Population Ecology 24: 174-192.
[7.] Wilson, Edward O. 1995. Naturalist.
[8.] Wilson (1975), 3.
[9.] Lewontin, R. C. (1979) `Sociobiology as an adaptationist program.'
Behavioral Science 24: 5-
14.
[10.] Ghiselin, Michael. (1973) `Darwin and evolutionary psychology.'
Science 179:964-968.
[11.] Pinker, Steven. (1997) How the Mind Works, 21.
[12.] Ibid., 34.
[13.] Mitchell, Melanie. "Can evolution explain how the mind works? A
review of the evolutionary
psychology debates." Santa Fe Institute.
[14.] Daly, M., & Wilson, M. I. (1996) `Violence against step-children.'
Current Directions in
Psychological Science, 5, 77-81.
[15.] Sober, Elliott, and Wilson, David S. (1998) Unto Others : The
Evolution and Psychology of
Unselfish Behavior.