Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr....

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Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of Christian Philosophy Karl-Rahner-Platz 1 A-6020 Innsbruck, Austria [email protected] www.uibk.ac.at/philtheol/loeffler 1. Aims & method of this lecture 2. The received view: Aristotle’s biology 3. Evolutionary ideas until Darwin 4. History of evolutionary thought from Darwin to the present 5. Evolution: Theory-structures and concepts 6. Evolution between science and world-view 7. Summary & discussion

Transcript of Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr....

Page 1: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015

• Prof.Dr.Dr. Winfried Löffler

• University of Innsbruck• Department of Christian

Philosophy• Karl-Rahner-Platz 1• A-6020 Innsbruck, Austria• [email protected]• www.uibk.ac.at/philtheol/loeffler

1. Aims & method of this lecture

2. The received view: Aristotle’s biology

3. Evolutionary ideas until Darwin

4. History of evolutionary thought from Darwin to the present

5. Evolution: Theory-structures and concepts

6. Evolution between science and world-view

7. Summary & discussion

Page 2: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

1. Aims and method of this lecture

Aims:

Some basics about current evolutionary biology

Some basics about its historical backgrounds

Understand its peculiar theoretical status

Distinguish between scientific theory and its ideological interpretations /

reductionisms

Misunderstandings of anti-evolutionism

Method: historical (2-4, partly 6) to understand conceptual backgrounds

Page 3: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

2. The received view: Aristotle’s biology

2.1 Life and Works

Aristotle (384-322) today usually seen as philosopher, but first big encyclopedist

Almost all scientific disciplines: - Physics, biology, physiology, meteorology, psychology, - economics, politology, aesthetics / poetology

- ethics, rhetorics, philosophy of language, - philosophy of science- general philosophy (“first philosophy”, metaphysics)

Page 4: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

Influential merits in the philosophy of science:

- Important basic concepts (substance/accident, potential/actual, matter/form, efficient cause/final cause)

- Classification of the scientific disciplines (theoretical/practical/poietical); (episteme / historia / techne)

- Structure of scientific arguments (Prior Analytics, syllogism)

- Structure of scientific classifications

- Structure of an empirical science (empirical “first sentences” & logical derivations from them) – overcoming Platonism

Heavy influence on Western thought!!!

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2.2 Biological writings

- Historia animalium (History of Animals) - see later- De partibus animalium (Parts of Animals)- De generatione animalium (Generation of Animals)

- Smaller writings (On movement of animals, sleep and sleeplessness, breathing, life and death etc.)

- Biggest part of his work is natural science!

- Interestingly: little interest in medicine (son of a healer!) and botany

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2.3 Aristotle’s method in biology

Empirical (broad sense) with speculative assumptions

Classification of phenomena plus question for causes (“4 causes”)

Reports of “experts”, own observations

Maybe: “experiments”, anatomical sections (stages of fertilized eggs)

Only scarcely: quantification (lengths); not weights, food intake etc.Only scarcely: ecological view (what it eats)

Aim: Collection/classification of facts (historía) and research for causes

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2.3 Aristotle’s method in biology

Teleology without design:

Natural things seen as functional units with functional parts

Teleological world-picture:

Chief pattern of explanation: final explanation, what is it good for?

Example: why do animals with a lung also have a neck? De part.An.III,3

(Roughly): Bipartite lungs need some tube to partition the air; that tube needs a certain length, hence the windpipe. Hence also, the oesophagus/jednjak. Hence, the necessity of a neck. (Fish don’t need one). The vicinity of windpipe and oesophagus is technically bad and would cause trouble; hence, nature contrived the epiglottis.

Nature as a whole is rational. Natura nihil facit frustra. Form follows functionBUT: No external “design plans”.

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2.4 Aristotle’s taxonomy

• 550 species according to morphological criteria, 300 re-identifiable today

• Groups and similarity observations: e.g., “all live-bearing quadrupeds have lungs and windpipes”

• Attempts to a classification of the whole range of animals, morphological features; not always consistent and sometimes wrong

• Man is included as a sub-class!

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2.4 Aristotle’s taxonomy

An example from HA IV 1, 524a3-20: “The octopus uses its tentacles both as feet and as hands; it draws in its food with the two that are placed over its mouth. The last of them, which is very sharp and is the only one which is whitish in colour and bifurcated at the tip—it is made so as to uncoil on the rhachis side (the rhachis being the smooth surface of the tentacle away from the suckers)—this one it uses in the act of copulation. In front of the sac and above the tentacles they have a hollow tube, by means of which they discharge from the sac any sea-water which may have come in while taking food into the mouth. The animal can move this tube to right and to left; it also discharges its “ink” through it. It swims obliquely in the direction of the so-called head, stretching out its feet; and by swimming in this way it can see forwards (since its eyes are on top), while its mouth is at the rear. So long as the animal is alive, the head is hard and as it were inflated. It takes hold of things and retains them with the underside of its tentacles, and the membrane between its feet is kept extended in its entirety. If it gets on to the sand, it can no longer retain its hold. […]”

Page 10: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

Aristotle’s taxonomy (roughly):

Animals with blood: (today: vertebrates)Live-bearing quadrupedsEgg-laying QuadrupedsBirdsFishCetacea (sea mammals)Egg-laying footless (snakes)Live-bearing footless (vipers)Man

Animals without blood:CephalopodsMolluscsInsectsCrustaceans

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2.5 The stability of species

Fundamental for 2300 years.

Minimal traces of evolutionary ideas?

(1) “Libya always produces something new” (De gen.an. II,7):

crossing of animal species at waterholes; reflection about the infertility of hybrids

(2) Reflection about Empedocles’ (5.century BC) mythical explanation of the origin of life: plants first, then parts of animals, then animals, only the useful ones survived

Aristotle: random combinations would not have survived (Phys. II, 8)

(3) Knowledge about variation: not all animals conform to species, monstrosities,

freak animals etc. (Not worrifying, just “freaks of nature”)

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2.6 Astonishing pioneerhoods and errors

Pioneerhoods:Sexual life of octopusOpposition to preformation theory / homunculus theory of the sperm (until 19th

century!). “Epigenetic”, step by step, formation of organs – the important ones for the genus-membership first.

Errors:Bison throws feces 7m for defenseWomen have less teethDayfly has only 4 legs Speculations:Parthenogenesis and other forms of reproductionProcreation (in mud etc.) – until Pasteur, 19th century! Brains serve to cool bloodWomen are incomplete men (more influence of katameria than of sperm)Women get soul later, birth rates m/f have to do with winds

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3. Evolutionary ideas until Darwin

3.1 Some aspects of biology after the middle ages

Renaissance: • Excursions, geograph. discoveries, herbaria, museums• flourishing of anatomy (Andreas Vesalius, Fabrica

(1543; De humani corporis fabrica libri VIII); public section of corpses; previously unknown exactness; descent-line: apes – “pygmies” (Plinius!) – man)

17th cent.: • new science of nature (Bacon, Galilei)

Experiments (planned variation of conditions, protocols) • New tools of observation: microscope, telescope; discovery

of micro-organisms, fine structures, insect development• “Physico-theology” & speculative preformationism (1695)

(hot discussions 18th cent. (Spallanzani), end 1830!)

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3.2 Carl von Linné’s taxonomy

• Various attempts to classifications, morphology comes in focus (microscopes!)

• Tension: Theory still Aristotelian (tree / pyramid), experienceshows multiplicity and similarities across branches

• “Natural history” becomes a discipline of its own

Carl von Linné (Linnaeus), 1707-1778; Swedish medic and botanistReform of taxonomy, 3 merits:a) New system of plants, with classification method (according to

number and structure of reproduction organs; “sexual system”)Systema naturae 1735; Genera plantarum 1737 etc.

b) Binary nomenclature (instead of descriptions), e.g. Sambucus nigra (sambucus = genus, sambucus nigra = species)

c) Terminology for the parts of plants

Page 15: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

3.2 Carl von Linné’s taxonomy

Backgrounds of Linné’s thought:

Objective structures in nature, ideas of the divine creator.

Not Aristotelian essentialism, but rather Enlightenment’s ideal of ordering

Does classificatory relatedness imply anything about historical relatedness?

Does the Systema point to a relation?

- Linné understood it as an artificial system with the task of ordering/quick finding; “natural system” as final project at horizon. Growing discontent about Linné´s artificiality.

- Some unclear remarks about an origin of plants

- Controversial how firmly he believed in the stability of species

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3.3 Early French speculations about evolution Georges Buffon (Histoire Naturelle, 36 vols., 1753-1788):

• Complexity and similarities in nature make a Linnéan classification impossible. And: “species” is an abstractum; there are individuals

• Procreation and change, by climate etc. Related species might have common ancestors, maybe one. Earth out of a collision sun-comet.

• Evolutionary scale/ladder instead of Linné´s hierarchical classification

• Evidence: similar anatomy across many species (donkey/horse, man/ape, man’s foot / horse’s foot etc.); rudimentary, useless organs

• Indirect message: EITHER God made the species by variation of few plans (great!), OR they have a common history. (Officially, the first…)

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3.3 Early French speculations about evolution Georges Cuvier (1769-1832), Étienne Geoffroy Saint-Hilaire (1772-

1844) and the Paris Academy Dispute 1831: Idealist versus evolutionary morphology?

Cuvier: “King of functional anatomy”, fine drawings, function determines structure, structure allows conjecture to function. “Reconstruction of died-out animals out of a few bones.”

Four basic construction plans of animals

Yet firm opposition to any evolutionary change (cats from Egypt, …)

Fossils are just extinct species, in global/regional catastrophes.

No “intermediate forms” in fossils

Geoffroy St.Hilaire: Research in homologies (e.g. gill bones in fish – ear bones in humans; vertebrate is similar to inverted worm (nerves at back, intestinal at front). Hence, change of species, common ancestry.

Political relevance: (1) not ideas of God (2) ...if not even nature is stable?

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3.4 Romanticist Philosophy of Nature and Morphology Johann W. von Goethe (1749-1832): • “morphology” instead of “comparative anatomy”: plants

and animals represent certain ideal shapes/forms/con-struction plans (mostly superficial, restricted to outer form)

• speculations about construction plan of a “Urpflanze” (ideal plant) which is realized in variants in real plants

• Postulates that skull bones are modified rib bones• Discovers the intermaxillary bone in human embryos: since humans

represent general mammal plan, and all other mammals have it, it must be somewhere… - in embryonal development!

“Göttingen School” (Carl Friedrich Kielmeyer, Johann Friedrich Meckel, Lorenz Oken, influenced by Fichte and Schelling): “Recapitulation”: Development of the embryo recapitulates animals of lower complexity

• Partly empirical (embryology, anatomy), partly philosophical: “formative power” of nature etc.; idealist “great chain of beings”

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3.4 Romanticist Philosophy of Nature and Morphology

Richard Owen (1848): Theory of “archetypes” (ideal design plans). Background: similar organs in very different animals (mole’s hand & dolphin’s fin have same bones!) cannot be due to environment.

Archetype of all vertebrates:

Terminological proposal, used till today:

Homologous organ: same construction, different function (e.g. mole’s hand, dolphin’s fin)

Analogous organ: different construction, same function (e.g. bird’s and butterfly’s wing)

Page 20: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

3.5 The first “evolutionary theory”: Lamarck 1809

Jean Baptiste de Lamarck (1744-1829): Philosophie zoologique (1809)

• “Lamarckism” today: inheritance of individually acquired characteristics, usually seen as overcome since Darwin; partial revival today

• First evolutionary theory: a theory that & how species change in the course of time.

• Simple forms of life emerge constantly from anorganic matter, procreation; hence, no common ancestry

• Inner tendency to higher development, “complexifying force”: le pouvoir de la vie

• Organisms adust behaviour to environment and inner state• Use/disuse of organs leads to growth/change of organs (e.g. neck)• … and that change is inherited! • As with Geoffroy St.Hilaire later: politically dangerous ideas…

Page 21: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

3.5 The first “evolutionary theory”: Lamarck 1809

Jean Baptiste de Lamarck (1744-1829): Philosophie zoologique (1809)

– “First Law: In every animal which has not passed the limit of its development, a more frequent and continuous use of any organ gradually strengthens, develops and enlarges that organ, and gives it a power proportional to the length of time it has been so used; while the permanent disuse of any organ imperceptibly weakens and deteriorates it, and progressively diminishes its functional capacity, until it finally disappears.”

– “Second Law: All the acquisitions or losses wrought by nature on individuals, through the influence of the environment in which their race has long been placed, and hence through the influence of the predominant use or permanent disuse of any organ; all these are preserved by reproduction to the new individuals which arise, provided that the acquired modifications are common to both sexes, or at least to the individuals which produce the young.”

Page 22: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

3.6 Erasmus Darwin (1731-1802) and a widely forgotten Anonymous: Robert Chambers (1844)

Erasmus Darwin (grandfather of Charles Darwin and Francis Galton)

Speculative views about Lamarck-style evolution:

“Would it be too bold to imagine, that in the great length of time, since the earth began to exist, perhaps millions of ages before the commencement of the history of mankind, would it be too bold to imagine, that all warm-blooded animals have arisen from one living filament, which the great first cause endued with animality, with the power of acquiring new parts, attended with new propensities, directed by irritations, sensations, volitions, and associations; and thus possessing the faculty of continuing to improve by its own inherent activity, and of delivering down those improvements by generation to its posterity, world without end!”

Glimpse of natural selection

[R. Chambers] Vestiges of the Natural History of Creation (1844)Popular science book, rather speculative, many errors

– Author: Robert Chambers, Scotch writer, anonymous

– Variability of species; Man descends from simpler forms of life

– Popularity and (slowly starting) excitement

Page 23: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

4. History of evolutionary thought from Darwin to the present

4.1 Charles Darwin (1809-1882) – life and worksSon of a religion-skeptic physician & unitarianStudies briefly medicine, natural history, theology; future: parson?No professional education as a scientist (biology not yet established!)

Invitation to travel on mapping-ship Beagle 1831-36Collects, studies, draws, sends

samples homeInitially only few doubts about

stability of speciesInfluences:Charles Lyell (geologist, big changes

of the earth surface; but rejectsLamarckism; species are created;they die out due to external change, and suppression by other species)

Richard Owen and RomanticsErasmus Darwin Lamarck (via Robert E. Grant!)

Page 24: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

Increasing doubts in Lyell’s biology (not in his geology!):

• released domestic animals adapt quickly to new environment • two similar ostrich species in overlapping areas (why does the one

not repress the other one?)• why do species dies out without change in environment? • why are died-out mammals replaced by other, similar ones?• the finches from the Galapagos islands:

- related with each other, related with animals on the continent, - different birds in similar habitats!- No clear-cut line “species” - “variety”

Page 25: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

Back in Cambridge

• Orders his findings, discusses (with Lyell, Owen, J. Gould (finches!))…

• Early speculations about an explanation for relations

• 1837: first sketch “I think”, insight that species must be changeable

Darwin’s speculations still +/- Lamarckist: - changes/variations are always useful, since- adaptation to environment as an embryo, geograph. isolation may cause new species - changes are inheritable

• 1838: reads Thomas Malthus: Essay on the Principle of PopulationPopulations have tendency to grow infinitely, but limited resources cause concurrence and limit growth,

Darwin: Selection not as embryo, but after birth. Theory of natural selection was +/- finished by 1839 (diaries).

Page 26: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

The Theory of Natural Selection

• Random, undirected, inheritable variations PLUS• natural selection under the pressure of the environment, concurrence for

food etc. … • … may in the long run lead to the emergence of adaptive (=useful)

change and new species.

• I.e., variations need not be perfect anymore. Old idea of perfect adaptation (from natural theology!) abandoned. Variations need just be slightly better.

• A manuscript was finished by 1842. Why not published? (1) Fear of scandal (Vestiges 1844!)? (2) Darwin had not yet a good idea for the ramifications / divergences in the changes.

• After 1854: Sympatry (different species in same area) is more important than Allopatry (different species in isolated areas). Coherent with the facts. Reason: Big areas have more ecological niches!

Page 27: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

The Publication and its effects

1856 Darwin presents his ideas to Lyell etc., friends urge to publish

1858 Alfred Russel Wallace sends a manuscript to Darwin, similar ideas

1858 Common paper by Darwin and Wallace, Wallace admits priority

1859 On the Origin of Species by Means of Natural Selection

Public effect:

In England dampened by a theol. discussion aboutcorrect Bible interpretation 1860, and Vestiges

John Herschel (astronomer, 1792-1871): unscien-tific, since only statistical

Thomas Huxley (“Darwin’s Bulldog”) tries to elicitcultural discussion, free universities, no religion

Not even whole church opposes. Still: excitement.

In France Cuvier dominates, no big interest

In Germany: bestseller, quick translation, popularizedby Ernst Haeckel (materialist reading!)

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Later Publications

1871 The Descent of Man (no big scandal; evolution of moral behavior)

1872 The Expression of Emotions in Animals and Man

Works about coral reefs,

plant fertilisation, rainworms, etc.

Darwin and religion:

1851 Death of his daughter, loss of religious faith, respectful agnostic

1882 Tomb in Westminster Abbey

Page 29: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

4.2 The Five Basic Tenets of Darwin’s Darwinism (E. Mayr)

• Evolution: species come and go through time, and while they exist they change. (But: the real bearer of evolution is the individual, not the species!)

• Common descent• Species multiply: the diversification of life involves populations of one

species diverging until they become separate species• Gradualism: evolutionary change occurs through incremental small

steps; new species are not created suddenly.• Natural selection: some variants change individual’s survival &

reproduction probability.

• Against Lyell: species are not stable; not perfectly adapted to environment; species are not created quickly (Lyell was a theory of species sequence, not of species evolution!)

• Against Lamarck: individual changes are not inherited; common descent

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4.3 Darwin’s Speculations about the biological fundament

• Notabene: Darwin had no idea about the place and nature of genetic information, laws of inheritance (Mendel’s rules etc.)

• Theory of Pangenesis: all cells produce little “gemmulae” (little buds), they gather in reproduction organs. In the young they mix

• = a Lamarckist remainder in Darwin!

• Still even in 1868 (book on selective breeding of animals and plants). Soft inheritance, acquired traits are passed on.

Page 31: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

4.4 Some open questions in Darwin

• Small steps or jumps? (Is mutation or selection the more important driving force of evolution?)

• Why sudden speeding up of evolutionary change, rapid growth of species numbers in certain stages of earth history (“Cambrian explosion” etc.)

• No real theory about inheritance, similarity and variability

• Why is there life at all? No procreation (no conflict with Pasteur 1859); maybe 4 or 5 original forms of life. Reference to creator more courtesy?

• Speculation about the warm little pond (letter to Hooker 1871):“But if (and Oh! What a big if!) we could conceive in some warm little pond, with all sorts of ammonia and phosphoric salts, light, heat, electricity etc, present, that a protein compound was chemically formed ready to undergo still more complex changes...”

• Similarity to Miller-Urey experiment 1953, “primordial soup” …

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The Miller-Urey experiment

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4.5 Ernst Haeckel (1834-1919): Materialism with idealist traces

• Great graphic illustrator, popularizer in Germany• Materialist (Darwinism as anti-religious and anti-conservative program!)

• Yet: some idealist remainders, idea of dynamics of growing complexity:– takes on recapitulation-idea of Kielmeyer

– Creates terms, and “biogenetic fundamental law”: “ontogeny” repeats “phylogeny”

– Evolutionary morphology: similarity of gastrula stage; speculation to a “Gastrea” as common ancestor

“tree of life” suggests height

of development, dynamics to complexity. (false: successful simple organisms, backward-developments)

Page 34: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

4.6 Francis Galton (1822-1911): Evolution becomes statistical

• Cousin of Darwin, multidisciplinary scientist: statistics, fingerprints, efficacy of prayer, questionnaire, blood-transfusion, eugenics

• Experiments against Darwin’s “gemmulae”• semi-lamarckian speculative theory of inheritance

• Important: Separation Organism – genetic information (“stirp”). Organism is just a representative selection from stirp, like a parliament

• Inheritance becomes statistical matter, every parent contributes ½

• Variation is now a matter of the population (not the organism, not the species!). New subject of evolution. Farewell to Aristotelian ideas.

• Beginning of theoretical biology as a mathematicised discipline!

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4.7 August Weismann (1834-1914): The organism as mere vehicle of genetic information

• Radical materialist• Similar to Galton, but with more empirical justification: experiments

with sea urchins (ježinac): germ cells are separated from the rest of the organism in very early stage

Separation germ cells – somatic cells

• Half-speculative theory of “immortal germ plasm”.

• Revolutionary: genetic info isthe bearer of variation, notthe organism; only “vehicle”

• Separation: growth/reproduct.• “Lamarck finally dead”• Cf. Dawkins’ Selfish Gene !!

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4.8 Gregor Mendel (1822-1884) The first robust theory of inheritance

OSA monk and abbot in Brno (CZ). Experiments with peas etc., publication 1865 in Austrian Journal, no reception. Only from 1900!

Question in those times: Does evolution go uniformly or in jumps?

Darwin: (Newton’s ideal! Natura non facit saltus!): uniformly

Huxley, Galton: undirected, small mutations / variations

BUT: such undirected small changes can cause no big changes, since statistics equalizes their effect. Statistical samples tend to average.

Hence, there must be other factors, Mutation must be more important.

Mendel’s method: reduction to only few (Y/N, not gradual) features, quantification (counting)

Experiments with crossing of peas and beans; features: white/lilac flowers; white/red/pink flowers, smooth/wrinkled seed.

Result: Mendel’s rules

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4.8 Gregor Mendel (1822-1884) The first robust theory of inheritance

Mendel’s first law (+ some more)

Presumptions: unmixable, discrete factors of inheritance

mutation more important than selection, bigger jumps are possible

But: biometry, statistics show continuous slow change. How can this

possibly be harmonized? “Great crisis of Darwinism” in early 20th century!

Page 38: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

4.9 The Molecular Revolution in Evolutionary Biology

End of 19th century: discovery of chromosomes in cell nucleus

1903/04 Sutton/Boveri: genetic info stored in chromosomes

1910 Johannsen: Terminology “Genotype” (inner constitution, info)

“Phenotype” (appearance);

Genotype Phenotype!

The genotype is inherited, not directly the features!!

After 1909 Drosophila experiments (quick reproduction, only 4x2 chromos.)

Beginning of localisation of genetic info on chromosomes (chemistry still unknown till 1953!)

Insights: - some features controlled by more than one gene

- there are interactions between genes

- one gene can influence more than one feature

HENCE: 1:1-matching “genes features” false since early 20th century!

“genetic blueprint” metaphor likewise wrong (still it is around…)

Page 39: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

4.9 The Molecular Revolution in Evolutionary Biology

1953 Watson/Crick: Discovery of double-helix structure of desoxyribonucleic acid (DNA)

Since mid-1960: Mechanisms of gene expression (DNA – RNA – proteine synthesis) begin to be clarified

“Human genome project” 1990-2003

Mind the correct way of speaking:

“Sequencing the genome”.

(Not: “decoding the genetic code”

– fallback to blueprint model!)

Page 40: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

4.10 The “Modern Synthesis” (1920s, 1930s onwards)

Darwinism had no undisturbed victorious career; no uniform development of post-Darwinist biology. Especially Mendel’s genetic discoveries caused worries: quick changes, “either/or” are possible. Mutation is more important!

Darwin’s intuition: small, incremental steps, selection is more important

“The great crisis of Darwinism” / “Eclipse of Darwinism” (J.Huxley 1942)

Late 19th/ early 20th century: all 5 tenets of Darwin come under doubt, many biologists step back to older theories, partly speculative.

Extension of population genetics (Ronald Fisher, Sewall Wright, John B. S. Haldane): Complex mathematical models to combine Mendel & Darwin

Some ideas from population genetics:• On genotype level Mendelian jumps, on phenotype level only small

Darwinian changes: since features often depend on many genes!• Population (not individuals) as the unit of evolution, statistics equalize:

in some individuals Mendelian jumps, in population slow Darwinian shift• The smaller a population, the easier are big jumps possible.

Page 41: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

4.10 The “Modern Synthesis” %

Theodosius Dobzhansky, Ernst Mayr and others:

“The Modern Synthesis”, “Synthetic Biology of Evolution”…

…connects: Darwinian core idea (stepwise evolution, selection)

Mendelian genetics

Population genetics

Biometry

Microbiology

Biochemistry

Behavioral science

Paleontology

Geology

etc.

Page 42: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

4.11 Some special ingredients of the Modern Synthesis

Genetic drift / bottleneck / founder effect

(A non-adaptive effect, against pan-selectionism!)

Punctuated equilibrium (Eldredge/Gould)

Various explanations of stages of stasis

and rapid evolutionary change; allopatric

origin of species (related with genetic drift)

Neutralism (Kimura): many big changes in DNA come and go,

without being tested by selection, neutral changes possible

%

Page 43: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

4.11 Some special ingredients of the Modern Synthesis %

Cooperation between genes; “Regulator genes”, governing the expression of other genes, “genetic switches”. E.g. Hox genes, regulate development of legs, antennas etc. mutations in such genes may have big effects!

“Evo-Devo” (Evolutionary developmental biology): • similarities in the genotype are astonishingly high across species, but • phenotypes differ dramatically!• Hence: genes in themselves are not so important• Hence: difference must lie in the expression of the genes,

development of the organism, regulator genes etc.• Evolutionary biology in the past took adult organism (and populations

out of them) as subject to natural selection,• EvoDevo takes the whole process/cycle of development of an

organism as the subject of natural selection.

Page 44: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

5.Evolution: Theory-structures and concepts

5.1 Theory-structure of EB

Physics, chemistry etc.: law-like explanations, able to prediction

false predictions a part of falsification

Evolutionary biology admits of (almost) no interesting predictions

Herschel: “un-scientific”, mere statistical relations

(young) Popper: unfalsifiable tenets like “the survival of the fittest”, circular.

Partial answer:

(1) Parts of explanations admit of predictions (cell level etc.)

Some few experiments with bacteria admit of prediction even of “direction of evolution” on macro-level: in lactose solution, bacteria with mutation develop genes to process lactose

(2) Why decreeing that all “scientific” explanation must resemble physics?

Page 45: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

5.2 Is EB a “science”?

Not a science like physics; combination of natural & historical disc.

Rather: a scientific research program, uniting many disciplines

Criteria by Philip Kitcher (The Advancement of Science, 1993)

A scientific practice… • Investigates an accepted domain of objects• Investigates accepted problems and questions• Has a non-natural technical terminology• Has commonly shared convictions at its basis• Applies accepted means and methods • Has accepted standards on aim and success of the investigations• Has accepted standards how to accepts results from other sciences• Is part of a social network

Evolution biology scores excellently under most criteria!

Page 46: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

5.3 The bearers / objects of evolution

Individuals / organisms?

Species?

Populations?

Genetic information?

Life-cycles? …different metaphysical views on objects of biology!

5.4 What are the traits / characteristics / features tested by evolution?

a) only bodily features (shape, food tolerances, …) or also behavior? Dawkins: “the extended phenotype” (including behavior) is being tested!

b) “Adaptive” traits (with a success story) – “maladaptive” traits – “by-products”

c) Is every feature adaptive? Panselectionism. (But: what is a feature??)

d) Beware of a frequent misunderstanding concerning “success story of trait”: - not the trait makes its success story (the trait is not there from beginning!) - rather: ex post, we see a success story of the trait and its previous traits (evolution does “bricolage” with available traits!). No “trait essentialism”

Page 47: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

5.5 Misunderstandings

“Survival of the fittest”: - Can be incremental; need not be perfect

- need not be “the strongest/biggest/…”

“Struggle for life”: - is not intra-population struggle

- need not be fight with similar species

- just general struggle with the ecological conditions: food, climate, safety, nesting-

places…

5.6 “Genes” (… an unsettled debate! “by the genes” is no explanation!)

“place on the chromosome”? – surely to imprecise

“sequence on the DNA” (concrete gene)? – things are more complicated: are “regulated /switched-on” genes really genes? Coding/non-coding DNA? Some DNA sequences are “read” more than once, some corres-pond to more than one proteine? Beginning/end sometimes unclear, …

“whatever makes a difference in fitness” (abstract, functional gene)? – may come close to antirealism concerning gene & genetic info.

Page 48: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

6. “Evolution” between science and world-view

6.1 Overview

SCIENCE:

Darwin’s (historical) Darwinism

“Darwinism” Synthetic theory of Evolution (1940 onwards)

Current EB (“New synthesis”, “EvoDevo”, …)

CONTROVERSIAL:

Pan-Selectionism

Sociobiology

“Cultural Evolution”, “Memetics”

NON-SCIENCE:

Social Darwinism

Pop-Darwinist Slogans

Page 49: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

6.2 Pan-Selectionism

Sometimes also: “strict neo-Darwinism”, “Darwinist orthodoxy”(Terminology seems not to be entirely fixed)

• Thesis: natural selection is the driving force of evolution; every feature has an “evolutionary success story” behind it

• Tendency to a deterministic account of the world and the human being: evolutionary successful features are +/- “hard-wired”, that includes also our mind, our reactions, etc.

• Critical evaluation: empirically unplausible in the light of modern genetics

• Ironically: a remainder of old “perfect adaptation” ideas (natural theology)

Page 50: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

6.3 Sociobiology

• Extension of evolutionary explanations also to our behavior, moral beliefs etc.

• First visions already in Darwin: “moral tribes” might be more successful

• Example1: why is altruism evolutionary successful?– At first glimpse: not at all. Risk your life, lose eating-time etc.

– Example “whistle-blowing” birds

– But kin-selection / group-selection (E.O. Wilson, Sociobiology 1975): brothers/sisters have 50% same genetic info, cousins 25%, … saving the life of, e.g., 5 cousins by whistle-blowing (and sacrifying own life!) increases the survival of own genetic info! 125 - 100= +25%

• Example 2: why are moral norms evolutionary useful?An economic way to secure cooperation of group members, internal/mental control instead of expensive external forces

• Critical evaluation: Empirically: unclear evidence• Manifold cultural shapings of behavior, limit natural/cultural unclear • Sociobiol. is no exclusive explanation (“altruism is nothing but xy…”)

Page 51: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

6.4 “Cultural evolution”, “memetics”

1976 Richard Dawkins, The Selfish Gene (and in later books)1995 Daniel Dennett, Darwin’s Dangerous Idea

• Basic idea: behavioral and cultural practices are passed on by imitation, they can be seen as items of “cultural information”: “memes”. Such memes undergo competition and selection, some do propagate, some do not

• Examples: termite-poking of chimpanzee tribes; language, use of fire, wheel, wall-graffiti, literary figures, cooking recipes, religion, product designs, business ideas…

• Critical evaluation: Suggestive examples…But huge differences between genes and memes:- genes mutate at random, memes often intentionally- genes from distant branches of evolution non mixable, memes are- memes are inherited in Lamarckist ways, individual inventions

In sum: misleading metaphor, parasitic on good image of biology. Even dangerous when mixed with ideas of neo-liberalism

Page 52: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

6.5 Social Darwinism

Label from 19th century, roots rather in Thomas Malthus and Herbert Spencer than in Darwin.

Basic ideas: (1) Evolutionary explains (and predicts) social facts(2) Evolutionary biology is not only descriptive (what is

the case?), but also normative (what should we do?)

(3) Evolution is progress

(4) There are “good” and “bad” genetic information-bits (for that progress!)

(5) Progress should not be retarded, “bad” genetic info should be suppressed (handicapped, lower races, …)

Usually: rivalry of all against all (people in a society, nations / nations, …), misunderstanding of “struggle for life”

Notabene: NS ideology was not closely connected with Darwin. Racism!

Page 53: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

6.5 Social Darwinism

Critical evaluation:

(1) Is/ought fallacy, description / normativity fallacy / genetic fallacy

(2) “Biological basis”of Social Darwinism is dubious and rather Lamarckist: Evolution is progress. But: progress/regress are not biological categories at all

(3) It depends on the environment which forms are “progressive / more successful” (small and undemanding animals might be most successful!)I.e., social Darwinism rests on some aesthetic ideals of what is healthy, strong, progressive etc.

(4) Even if we could create a “perfect natural world”, this does not guarantee a pleasant, humane culture. False genetic determinism, nature does not determine culture.

Page 54: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

6.6 Some pop-Darwinist slogans

“Nature and Evolution are nothing else but a huge process of chance and random”

– BIOLOGICALLY FALSE, at least selection is not random.

“Darwinism means that mutation and natural selection explain every feature.”

– BIOLOGICALLY MOST PROBABLY FALSE, only most narrow-minded pan-selectionists would deny other evolutionary effects

“Cosmic evolution”: “There is one big evolutionary process from Big Bang to human culture.”

– CONCEPTUAL NONSENSE. (1) Not every agglomeration and development process (as in the early universe) is also an evolution process. There is no multiplication, mutation, inheritance, and selection for protons, galaxies, stars etc. Over-extension of a biological metaphor. (2) On cultural evolution, see 6.4.

%

Page 55: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

6.6 Some pop-Darwinist slogans %

“Society and economy work according to evolutionary laws.”

– SEE 6.4 above!

“Science (like biology) consists of falsifiable claims only, world-views are irrational and subjective.”

– FALSE FROM THE VIEWPOINT OF PHIL. OF SCIENCE. (1) Too narrow concept of “science”

(2) Background assumptions are even at work in the sciences: see Kuhn’s paradigms, Lakatos’ “hard cores” of research programs. Border to world-views fuzzy; world-views contain rational structures, can be more/less plausible, and rationally discussable. See Popper: there are genuine philosophical problems, beyond science.

Page 56: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

6.7 Biology and Religion: On “Creationism”, “Intelligent Design” etc.

A bit of history

USA a country of small religious groups, partly literal reading of the Bible

About one half of the population has serious doubts about evolution

Empirical fundament: unclear spots in evolution, cosmic fine tuning

Spectrum:• Short-time (Young-earth) creationism 6 days, about 6000 yrs ago• Long-time creationism (special creationism) successive intervention• “Intelligent Design” design-plans, to solve irreducible complexity• “Theistic Evolution” (Vatican II; God as 1st cause carries 2nd causes)• Deism• Atheism, metaphysical naturalism

Wealthy think-tanks, www.discovery.org etc., disinformation, wedge

Page 57: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

6.7 Biology and Religion: On “Creationism”, “Intelligent Design” etc.

Objections to ID

• Unclear ontology of “design plan”, at odds with usual scientific ontology• Selective evidence, focused on explanatory gaps and failures of EB• Some irreducible complexities are reducible; regulator-genes etc. may

explain complexities • “Filling-the-gap-theology”: every new biological discovery makes God

more irrelevant, theology on constant retreat• ID proves only world-constructor (demiurg), not creator• Why a single God behind the 1000’s of appearances of ID? Tacit

background theology smuggled in.• Problem of evil radicalized: source of bad design, senseless design?

• In sum: neither necessary nor fruitful for the Christian.• (Due!) criticism of reductionism and ideology should not turn into (undue)

pseudo-scientific blends of theology and biology.

Page 58: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

7. Summary

• Aristotle inaugurated a vision of biology with the species in focus, and a teleological view at the world – yet, without external design.

• The raise of modern evolutionary thought began before Darwin and was based on empirical findings as well as on speculative ideas. Darwin and Wallace developed the first coherent, large-scope theory with a broad empirical basis (still with many blind spots).

• The basic ideas of Darwin are still topical in the core of current EB. However, EB can’t be reduced to “mutation and selection”.

• Current EB is a research program with many open questions, especially about the impact of natural selection. Central concepts like “gene” admit of different readings.

• This is one reason why EB has a strong affinity to be reduced to an ideology, more than other sciences. Another reason is the wide scope of EB, from the explanation of shape to behavior.

• ID is a dubious blend of theology and biology which appears neither necessary nor useful for the theologian.

Page 59: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

8. Repetition questions

1. How does Aristotle’s conception of science differ from Plato’s?2. Name 4 of Aristotle’s biological writings and their (rough)

content.3. How does Aristotle’s method in biology resemble/differ from

today?4. What type of discipline is an episteme, what a historia? 5. Why did we describe Aristotle’s biology as „teleology without

design“?6. Sketch Empedocles’s explanation of the origins of life and

Aristotle’s counter-argument.7. Aristotle’s doctrine of the stability of species (what is it?) is said

to have prevented evolutionary ideas for centuries. But are there any traces of variability of species in Aristotle?

8. How did Aristotle describe the development of an embryo?9. What is preformationism?10.What is procreationism?11.What was new about Linné’s taxonomy system? What are its

limits? Was he an Aristotelian?

Page 60: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

12. What other inventions did he make which are still important for biology?

13. Sketch Buffon’s anti-Linnéan position.

14. What was the Paris Academy Dispute 1831 ultimately about? (Don’t just say “idealist versus evolutionary morphology”, but explain the issue!)

15. What are “homologous” organs, what are “analogous” ones?

16. Explain some of the basic intuitions of Romanticist biologists, and name some important persons.

17. Why can we say that Lamarck presented the first evolution theory? Only under what conditions should something be called an evolution theory?

18. What does crucially distinguish “Lamarckism” from standard evolutionary biology?

19. How did Lamarck explain the change of organisms in the course of time?

20. Name some thinkers and researchers who influenced Darwin!

21. How did Malthus change Darwin’s thought?

22. Why is Darwin’s theory of natural selection no theory of perfect adaptation?

23. What is sympatric, what allopatric origin of new species?

24. How did Ernst Mayr summarize classical Darwinism?

Page 61: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

25. Sketch Darwin’s “gemmulae” theory. What was it good for, what was it supposed to explain?

26. List some open problems in Darwin.27. Why are Haeckel’s tree-diagrams suggestive in a false

direction? 28. What does his “biogenetic fundamental law” say? 29. Describe the separation of genetic information from the

organism in Galton, Weismann and Dawkins. In what are stirp, germ plasm and selfish gene similar?

30. What new point of view dis Galton bring into biology? Why does this bring a turn away from Aristotelian ideas?

31. Why was the “Weismann barrier” seen as a death-blow to Lamarckism?

32. Why did Mendel’s discoveries shake the Darwinism of his time? What was the worrisome problem?

33. Sketch some main steps in the discovery of genetic information.34. Why is “we can today decode the genetic code” a problematic

way of speaking?35. Why can population genetics perhaps unite Darwinist and

Mendelian ideas?36. What does “pan-selectionism” claim?

Page 62: Philosophical presuppositions of evolutionary biology FFDI Zagreb, 20-25 April 2015 Prof.Dr.Dr. Winfried Löffler University of Innsbruck Department of.

37. Why is the (empirically backed!) idea of genetic drift an argument against pan-selectionism?

38. Try to describe some basic ideas of EvoDevo. Why is EvoDevo a remedy against the misconception of a “genetic blueprint”?

39. How do evolutionary explanations differ from physical explanations?

40. Why has evolutionary biology a special structure, different from other natural sciences?

41. What units/bearers of evolution were proposed in the history of biology?

42. Why is it misguided to say “feature F had an evolutionary success story”?

43. What did Darwin mean by “struggle for life”?44. Sketch different notions of a “gene”.45. How would socio-biologists explain that altruism is sometimes

evolutionary successful, even if it goes to sacrificing one’s life?46. What is a “meme”? Why is “cultural evolution” a non-starter?47. Describe the mistakes of “Social Darwinism”.48. Why is “Intelligent Design” not a wise choice for theologians? 

GOOD LUCK FOR THE EXAM!