Lessons from Slijper’s Goat:Lessons from Slijper’s Goat:On the Convergence of Classical and On the Convergence of Classical and

Modern BiologyModern Biology

James BarhamJames BarhamUniversity of Notre Dame andUniversity of Notre Dame and

Institute for the Study of NatureInstitute for the Study of NatureJune 12, 2008June 12, 2008

I. The Reality of Functional I. The Reality of Functional StabilityStability

Recall our definition of life: Recall our definition of life: Life is the Life is the inherent capacity of a certain kind of inherent capacity of a certain kind of material system for spontaneous activity material system for spontaneous activity resulting in the functional stability of the resulting in the functional stability of the systemsystem

Two questions:Two questions: Is “functional stability” real, or merely a kind of Is “functional stability” real, or merely a kind of

mental construct that human beings cannot help mental construct that human beings cannot help projecting onto living things?projecting onto living things?

If it is real, how is it physically possible?If it is real, how is it physically possible?

A. Slijper’s Goat as a Model A. Slijper’s Goat as a Model OrganismOrganism

Throughout the history of science, certain Throughout the history of science, certain organisms have come to exemplify organisms have come to exemplify general theories or attitudes toward life:general theories or attitudes toward life: 1818thth c.—Trembley’s “polyp” (freshwater hydra) c.—Trembley’s “polyp” (freshwater hydra)

(vitalism)(vitalism) 1919thth c.—Darwin’s finches (natural selection) c.—Darwin’s finches (natural selection) 2020thth c.—Morgan’s fruitflies ( c.—Morgan’s fruitflies (Drosophila Drosophila

melanogastermelanogaster) (genetics)) (genetics) 2121stst c.—Slijper’s goat (functional realism, c.—Slijper’s goat (functional realism,

emergentism)?emergentism)?

B. The Standard Reductionist B. The Standard Reductionist Account of FunctionAccount of Function

According to the standard account, the According to the standard account, the functional (teleological) aspects of living functional (teleological) aspects of living things can be fully reduced to physical things can be fully reduced to physical (“efficient”) causation, with the help of two (“efficient”) causation, with the help of two bodies of theory:bodies of theory: Cybernetic controlCybernetic control Natural SelectionNatural Selection

Slijper’s goat is important because it is a Slijper’s goat is important because it is a vivid demonstration of the inadequacy of the vivid demonstration of the inadequacy of the standard accountstandard account

C. Story of Slijper’s GoatC. Story of Slijper’s Goat

E.J. Slijper (1907–1968) was a professor of E.J. Slijper (1907–1968) was a professor of veterinary anatomy at the University of Utrecht in veterinary anatomy at the University of Utrecht in the Netherlands (later Professor of Zoology at the the Netherlands (later Professor of Zoology at the University of Amstedam) University of Amstedam) (Moore 1969)(Moore 1969)

In 1939, Slijper received a three-month-old goat In 1939, Slijper received a three-month-old goat born without forelegs born without forelegs (Slijper 1942)(Slijper 1942)

Under his care, the goat learned to locomote by Under his care, the goat learned to locomote by means of an upright posture and kangaroo-like means of an upright posture and kangaroo-like bipedal gaitbipedal gait

Upon its death, autopsy revealed extensive Upon its death, autopsy revealed extensive remodeling of the muscles and skeleton, remodeling of the muscles and skeleton, resembling those of rodents and marsupials that resembling those of rodents and marsupials that normally adopt a bipedal gait.normally adopt a bipedal gait.

D. Faith the DogD. Faith the Dog

A contemporary case similar to A contemporary case similar to Slijper’s goat: Faith the DogSlijper’s goat: Faith the Dog http://www.youtube.com/watch?http://www.youtube.com/watch?

v=aZsV4R3XJKkv=aZsV4R3XJKk

E. The Undisputed Facts E. The Undisputed Facts Relating to Slijper’s GoatRelating to Slijper’s Goat

Locomotion is a crucial function in animals, normally Locomotion is a crucial function in animals, normally constructed soon after birthconstructed soon after birth

The construction of normal quadrupedal gait in the goat was The construction of normal quadrupedal gait in the goat was severely perturbed due to the birth defectseverely perturbed due to the birth defect

The organism of the goat compensated for this perturbation The organism of the goat compensated for this perturbation by spontaneously constructing the radically different, bipedal by spontaneously constructing the radically different, bipedal gaitgait

At a minimum, this construction of bipedal gait involved the At a minimum, this construction of bipedal gait involved the major restructuring of three organ systems:major restructuring of three organ systems: MusculatureMusculature SkeletonSkeleton Nervous systemNervous system

The construction of bipedal gait was the result both of The construction of bipedal gait was the result both of conscious striving (it required effort and learning), and also conscious striving (it required effort and learning), and also of non-conscious compensatory response throughout the of non-conscious compensatory response throughout the organismorganism

F. West-Eberhard’s F. West-Eberhard’s InterpretationInterpretation

West-Eberhard West-Eberhard (2005)(2005) invokes the case of invokes the case of Slijper’s goat with a view to revising our Slijper’s goat with a view to revising our understanding of evolutionunderstanding of evolution

The idea is that Slijper’s goat is a more The idea is that Slijper’s goat is a more extreme (and thus clearer) example of a extreme (and thus clearer) example of a normal and perfectly general phenomenonnormal and perfectly general phenomenon

Her claim is that all evolutionary change Her claim is that all evolutionary change begins as “phenotypic accommodation,” begins as “phenotypic accommodation,” meaning a compensatory response to meaning a compensatory response to perturbation, like that displayed by Slijper’s perturbation, like that displayed by Slijper’s goat, leading to an adaptive innovation, like goat, leading to an adaptive innovation, like the bipedal gaitthe bipedal gait

G. Phenotypic Accommodation G. Phenotypic Accommodation and Evolutionand Evolution

Phenotypic accommodation is made possible by Phenotypic accommodation is made possible by what she calls “developmental plasticity,” where what she calls “developmental plasticity,” where “development” is understood broadly to include “development” is understood broadly to include the ability of both the physical structure and the the ability of both the physical structure and the behavior of the organism to compensate for behavior of the organism to compensate for perturbation, both pre- and post-partumperturbation, both pre- and post-partum

Adaptive innovations constructed through the Adaptive innovations constructed through the normal process of phenotypic accommodation normal process of phenotypic accommodation then become genetically “fixed” (“genetic then become genetically “fixed” (“genetic accommodation”), leading to permanent accommodation”), leading to permanent evolutionary changeevolutionary change

The extent of genetic accommodation is highly The extent of genetic accommodation is highly controversialcontroversial

H. Conceptual Priority of H. Conceptual Priority of Phenotypic AccommodationPhenotypic Accommodation

Nevertheless, phenotypic accommodation should Nevertheless, phenotypic accommodation should not be controversialnot be controversial

Not only can we observe it in a case like Slijper’s Not only can we observe it in a case like Slijper’s goat, it also has conceptual priority within the goat, it also has conceptual priority within the framework of the theory of natural selectionframework of the theory of natural selection

A genetic change can only lead to a phenotypic A genetic change can only lead to a phenotypic change by the process of development, broadly change by the process of development, broadly construedconstrued

Adaptive innovations must first be Adaptive innovations must first be developmentally constructed before they can be developmentally constructed before they can be subjected to natural selectionsubjected to natural selection

Therefore, phenotypic accommodation is Therefore, phenotypic accommodation is conceptually prior to natural selectionconceptually prior to natural selection

I. Note on TerminologyI. Note on Terminology

A number of different terms are found in the literature to A number of different terms are found in the literature to refer to what West-Eberhard calls “phenotypic refer to what West-Eberhard calls “phenotypic accommodation”:accommodation”: PlasticityPlasticity RobustnessRobustness AdaptabilityAdaptability

Useful distinctions among these ideas can be made for Useful distinctions among these ideas can be made for various purposesvarious purposes

However, for our purposes, the main point is that when a However, for our purposes, the main point is that when a perturbation occurs within a living system, compensatory perturbation occurs within a living system, compensatory changes spontaneously occur in order to restore function changes spontaneously occur in order to restore function and preserve lifeand preserve life

This capacity for adaptive compensation is what we have This capacity for adaptive compensation is what we have been calling the “functional stability” of living thingsbeen calling the “functional stability” of living things

J. Our Interpretation: J. Our Interpretation: Functional Stability Is BasicFunctional Stability Is Basic

Slijper’s goat shows that functional stability is Slijper’s goat shows that functional stability is objectively real in the sense of being “kickable”: objectively real in the sense of being “kickable”: perturbations lead to observable compensationsperturbations lead to observable compensations

Furthermore, this case makes it obvious that this Furthermore, this case makes it obvious that this capacity cannot be explained on the basis of capacity cannot be explained on the basis of genesgenes

This example also helps to make it clear that far This example also helps to make it clear that far from being reduced by natural selection, from being reduced by natural selection, functional stability is presupposed by that theoryfunctional stability is presupposed by that theory

Not only is functional stability demonstrably real, Not only is functional stability demonstrably real, and not only is it unreduced—it is the foundation and not only is it unreduced—it is the foundation upon which evolutionary theory tacitly restsupon which evolutionary theory tacitly rests

II. Why Teleoreductionism II. Why Teleoreductionism FailsFails

These are some rather sweeping These are some rather sweeping claimsclaims

Let us now slow down a little bit and Let us now slow down a little bit and do some conceptual work with the do some conceptual work with the phenomenon of functional stability, phenomenon of functional stability, or what I shall now call “immanent or what I shall now call “immanent teleology,” and the claims of teleology,” and the claims of mainstream reductionismmainstream reductionism

A. Teleoelimination and A. Teleoelimination and TeleoreductionTeleoreduction

““Teleoelimination” is the claim that there Teleoelimination” is the claim that there is no such thing as immanent teleologyis no such thing as immanent teleology Flies in the face of both biological practice and Flies in the face of both biological practice and

common sensecommon sense ““Teleoreduction” is the claim that Teleoreduction” is the claim that

immanent teleology is “nothing but” immanent teleology is “nothing but” mechanistic processes of cybernetic mechanistic processes of cybernetic control put in place by natural selectioncontrol put in place by natural selection Somewhat more plausbileSomewhat more plausbile

B. The Concept of B. The Concept of TeleofunctionTeleofunction

The word “function” is used in various ways, so let The word “function” is used in various ways, so let us say “teleofunction” to be clear about our claim us say “teleofunction” to be clear about our claim that functional stability is a form of immanent that functional stability is a form of immanent teleologyteleology

Given a pair of events A and B with temporal Given a pair of events A and B with temporal ordering such that A = earlier (“cause”) and B = ordering such that A = earlier (“cause”) and B = later (“effect”), for a causal relation between A and later (“effect”), for a causal relation between A and B to count as a B to count as a teleofunctionteleofunction, there must exist a , there must exist a (partial) determination of B by A (A (partial) determination of B by A (A B), but B), but alsoalso a a (partial) determination of A by B (A(partial) determination of A by B (A B) B)

In this case, A = “means” and B = “end”In this case, A = “means” and B = “end” Absent such (partial) determination of the Absent such (partial) determination of the

temporally prior means by the temporally later end temporally prior means by the temporally later end (A(A B), there is no teleofunction (final causation). B), there is no teleofunction (final causation).

C. The Problem of Backwards C. The Problem of Backwards CausationCausation

All of life seems to operate according to the means-end All of life seems to operate according to the means-end logic of teleofunction; certainly, it is obvious that a very logic of teleofunction; certainly, it is obvious that a very great deal of our own human experience takes this great deal of our own human experience takes this formform

However, the standard account of biology denies that However, the standard account of biology denies that teleofunction is objectively real; it is considered at best teleofunction is objectively real; it is considered at best a necessary illusion due to human cognitive defectsa necessary illusion due to human cognitive defects

One great obstacle to accepting teleofunction at face One great obstacle to accepting teleofunction at face value is understanding how it could be possiblevalue is understanding how it could be possible

On the face of it, teleofunction seems to require a On the face of it, teleofunction seems to require a future state of affairs to exert causal influence upon a future state of affairs to exert causal influence upon a present state of affairs: But how could this be?present state of affairs: But how could this be?

This is the problem of so-called “backwards causation”This is the problem of so-called “backwards causation”

D. Representations Are No D. Representations Are No HelpHelp

One standard way of responding to the backwards One standard way of responding to the backwards causation problem is to invoke representationscausation problem is to invoke representations

The idea is that it is not a particular future state of the The idea is that it is not a particular future state of the world that influences the present, but rather a model world that influences the present, but rather a model or “representation” of that state that already existsor “representation” of that state that already exists

However, while this seems to work for animals with However, while this seems to work for animals with brains (which is where representations are supposed brains (which is where representations are supposed to be located), it does not help with the general to be located), it does not help with the general problem of functional stability, which is found in all problem of functional stability, which is found in all living thingsliving things

Moreover, it does not really work for animals with Moreover, it does not really work for animals with brains, either; let us see brains, either; let us see why why

E. The Problem of E. The Problem of NormativityNormativity

Representation is a Representation is a normativenormative concept; that is to say, concept; that is to say, it constitutes a it constitutes a criterioncriterion or or standardstandard according to according to which actions are judged right or wrongwhich actions are judged right or wrong

For example, to say that the word “cat” represents a For example, to say that the word “cat” represents a cat is to say that “cat” cat is to say that “cat” is supposedis supposed to mean a cat, and to mean a cat, and not some other animalnot some other animal

To utter the word “cat” (to an English speaker) while To utter the word “cat” (to an English speaker) while intending to represent a dog is to make a intending to represent a dog is to make a mistakemistake

But there is no such thing as success or failure in But there is no such thing as success or failure in physics and chemistry; matter just does what it must physics and chemistry; matter just does what it must do, it is never right or wrongdo, it is never right or wrong

Therefore, a representation is not a purely physical Therefore, a representation is not a purely physical thing; it already embodies an aspect of the problem of thing; it already embodies an aspect of the problem of teleofunction, and so cannot solve itteleofunction, and so cannot solve it

F. Goal, Norm, and ValueF. Goal, Norm, and Value

Another way of looking at teleofunction is to notice Another way of looking at teleofunction is to notice that a goal-state is a that a goal-state is a specialspecial or or preferredpreferred state of a state of a systemsystem

It is this specialness or preference that is the heart of It is this specialness or preference that is the heart of the problemthe problem

A preferred state of a system has A preferred state of a system has valuevalue for the for the system; that is, realizing the preferred state is system; that is, realizing the preferred state is goodgood for the system, and failing to do so is for the system, and failing to do so is badbad

It is the goodness of preferred states that underlies It is the goodness of preferred states that underlies the normativity of the means-end logic of the normativity of the means-end logic of teleofunction; the means are right insofar as the teleofunction; the means are right insofar as the state they produce (the end) is goodstate they produce (the end) is good

In short, teleology, normativity, and axiology are a In short, teleology, normativity, and axiology are a single problem-complexsingle problem-complex

G. The Failure of G. The Failure of Teleoreduction—Cybernetic Teleoreduction—Cybernetic

Control Control The two forms of teleoreduction—cybernetic The two forms of teleoreduction—cybernetic

control and natural selection—both fail for the control and natural selection—both fail for the same reason:same reason: They have no conceptual resources for explaining in non-They have no conceptual resources for explaining in non-

normative terms why one particular state of a system normative terms why one particular state of a system should be privileged over others as the preferred stateshould be privileged over others as the preferred state

The concept of cybernetic control presupposes an The concept of cybernetic control presupposes an external agent who determines what external agent who determines what countscounts as as the preferred state of the systemthe preferred state of the system The thermostat on my home heating system does not The thermostat on my home heating system does not

care if it is keeping the temperature of my house at the care if it is keeping the temperature of my house at the correct settingcorrect setting

It is I who determine what counts as the correct It is I who determine what counts as the correct temperature, in relation to which the thermostat is temperature, in relation to which the thermostat is judged as working properly or improperly judged as working properly or improperly

H. The Failure of H. The Failure of Teleoreduction—Natural Teleoreduction—Natural

Selection Selection The theory of natural selection likewise The theory of natural selection likewise

presupposes value and normativitypresupposes value and normativity It simply It simply assumesassumes that survival is the preferred state for that survival is the preferred state for

organisms organisms Means (traits) that are (relatively) successful at realizing Means (traits) that are (relatively) successful at realizing

their functional ends are the ones “selected”; it is not their functional ends are the ones “selected”; it is not “selection” that makes means capable of realizing their “selection” that makes means capable of realizing their endsends

Selection as such has no magic power to Selection as such has no magic power to transform a physical state into a normative transform a physical state into a normative preferred state, as shown by the Swampman preferred state, as shown by the Swampman thought experiment thought experiment (Davidson 2001)(Davidson 2001)

I. Backwards Causation I. Backwards Causation AgainAgain

But if teleoreduction fails, what is the solution?But if teleoreduction fails, what is the solution? The problem of backwards causation is a genuine The problem of backwards causation is a genuine

oneone After all, surely the future as such does not have any After all, surely the future as such does not have any

causal influence upon the presentcausal influence upon the present One solution is to see that “backwards causation” is One solution is to see that “backwards causation” is

ambiguous between:ambiguous between: (a) the idea that a non-actual state of a system can (a) the idea that a non-actual state of a system can

causally influence an actual state (which is essential to the causally influence an actual state (which is essential to the idea of teleofunction); andidea of teleofunction); and

(b) the idea that the future is in some sense actual (as on (b) the idea that the future is in some sense actual (as on certain “block-universe” interpretations of time), and by certain “block-universe” interpretations of time), and by virtue of this actuality may causally influence the pastvirtue of this actuality may causally influence the past

J. Preferred States as Virtual J. Preferred States as Virtual StatesStates

To solve the problem of backwards causation, we To solve the problem of backwards causation, we have merely to reject the idea that the future is have merely to reject the idea that the future is actual in any sense at allactual in any sense at all

This affirms the reality of the flow of time, which This affirms the reality of the flow of time, which is anyhow desirable on realist groundsis anyhow desirable on realist grounds

Since, on this interpretation, the non-actual state Since, on this interpretation, the non-actual state of a teleofunction (the end) that causally of a teleofunction (the end) that causally influences the actual state (the means) is not the influences the actual state (the means) is not the future as such, let us give it another name:future as such, let us give it another name: Let us call it a Let us call it a virtual statevirtual state

Therefore, according to this idea, Therefore, according to this idea, preferred states preferred states of teleofunctions are virtual statesof teleofunctions are virtual states

K. What Is a Virtual Preferred K. What Is a Virtual Preferred State of a Teleofunction? State of a Teleofunction?

The question is, then: Can we give the concept of The question is, then: Can we give the concept of a virtual preferred state of a teleofunction a a virtual preferred state of a teleofunction a plausible scientific interpretation?plausible scientific interpretation? The answer is YesThe answer is Yes Example: The notion of a high-dimensional phase-space Example: The notion of a high-dimensional phase-space

attractor incorporates a notion of virtuality that some attractor incorporates a notion of virtuality that some have suggested may be useful for modeling the have suggested may be useful for modeling the preferred state of teleofunctions (Delattre 1986; Yates preferred state of teleofunctions (Delattre 1986; Yates 1994) 1994)

In the final section, let us see how this idea can be In the final section, let us see how this idea can be put together with ideas of ontological emergence put together with ideas of ontological emergence to provide us with an alternative, realist view of to provide us with an alternative, realist view of the strange functional stability of lifethe strange functional stability of life

III. Toward a Teleorealist View III. Toward a Teleorealist View of Lifeof Life

The view of life I wish to sketch briefly The view of life I wish to sketch briefly here might be called “teleorealist”:here might be called “teleorealist”: That is, it views the immanent teleology That is, it views the immanent teleology

associated with the functional stability of associated with the functional stability of life as objectively reallife as objectively real

We shall see that the key to a We shall see that the key to a scientifically plausible teleorealism is scientifically plausible teleorealism is viewing the cell within the framework viewing the cell within the framework of ontological emergenceof ontological emergence

A. The “Solid State Cell”A. The “Solid State Cell”

The first step toward understanding how The first step toward understanding how functional stability can be physically possible is to functional stability can be physically possible is to realize that the cytoplasm is far from being in realize that the cytoplasm is far from being in aqueous phase aqueous phase (Luby-Phelps 2000; Walleczek 2000)(Luby-Phelps 2000; Walleczek 2000)

On the contrary, it is so tightly packed with On the contrary, it is so tightly packed with macromolecules that it has many properties in macromolecules that it has many properties in common with semi-solid phases of matter, such common with semi-solid phases of matter, such as:as: liquid crystals (liquid crystals (Ho et al. 1996)Ho et al. 1996); and; and gels gels (Pollack 2001)(Pollack 2001)

It has even been referred to recently as the “solid It has even been referred to recently as the “solid state cell” state cell” (McNiven 2003) (McNiven 2003)

B. Condensed-Matter Physics B. Condensed-Matter Physics of the Cellof the Cell

Though speculative, a number of field-Though speculative, a number of field-theoretic properties have been postulated theoretic properties have been postulated for cytoplasmic structures, including:for cytoplasmic structures, including: Long-range coherence via the so-called Long-range coherence via the so-called

“Fröhlich mechanism” “Fröhlich mechanism” (Del Giudice et al. 2005; Ho (Del Giudice et al. 2005; Ho 1997) 1997)

Long-range collective action via functionally Long-range collective action via functionally induced phase transitions induced phase transitions (Pollack 2001)(Pollack 2001)

““Quasi-particles” possibly governed by special Quasi-particles” possibly governed by special conservation principles conservation principles (Vitiello 2001)(Vitiello 2001)

C. How Does All This Help?C. How Does All This Help?

It is often pointed out that “self-organization” It is often pointed out that “self-organization” models that appeal to principles like dissipative models that appeal to principles like dissipative structures fail to distinguish living matter from structures fail to distinguish living matter from nonliving oscillators like hurricanesnonliving oscillators like hurricanes

Therefore, a natural question to ask is: How does Therefore, a natural question to ask is: How does condensed-matter physics help explain functional condensed-matter physics help explain functional stability?stability?

Tentative answer: Ontological emergence and Tentative answer: Ontological emergence and scale-transcendent theoretical principles allow us scale-transcendent theoretical principles allow us to envision emergent quantum numbers and to envision emergent quantum numbers and conservation principles specific to the living state conservation principles specific to the living state of matterof matter

D. The Master SpeculationD. The Master Speculation

We know that the functional stability of life cannot We know that the functional stability of life cannot be explained purely in terms of ordinary physical be explained purely in terms of ordinary physical principles such as minimum actionprinciples such as minimum action

But perhaps functional stability could be But perhaps functional stability could be understood in terms of some as-yet unknown understood in terms of some as-yet unknown conserved quantum numbers—say, corresponding conserved quantum numbers—say, corresponding to a “minimum effort principle” or to a “minimum effort principle” or “appropriateness” of means to end“appropriateness” of means to end Indeed, the faint outline of such an idea may already be Indeed, the faint outline of such an idea may already be

visible in notions like “minimum frustration principle” and visible in notions like “minimum frustration principle” and “topological charge conservation” that have been “topological charge conservation” that have been advanced in connection with models of the “functionally advanced in connection with models of the “functionally important motions” of proteins important motions” of proteins (Frauenfelder et al. 1991)(Frauenfelder et al. 1991)

E. ConclusionE. Conclusion

While this is all admittedly wildly While this is all admittedly wildly speculative at present, nevertheless I speculative at present, nevertheless I think it goes to show that a general think it goes to show that a general framework of ontological emergence framework of ontological emergence gives us some useful resources for gives us some useful resources for thinking about functional stability, thinking about functional stability, and for taking a teleorealist view of and for taking a teleorealist view of life, generallylife, generally

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