Physical explanations and biological explanations,

empirical laws and a priori laws

Joel Press
Received: 18 October 2006 / Accepted: 17 September 2007 / Published online: 25 October 2007
Springer Science+Business Media B.V. 2007
Abstract Philosophers intent upon characterizing the difference between physics and
biology often seize upon the purported fact that physical explanations conform more
closely to the covering law model than biological explanations. Central to this purported
difference is the role of laws of nature in the explanations of these two sciences. However,
I argue that, although certain important differences between physics and biology can be
highlighted by differences between physical and biological explanations, these differences
are not differences in the degree to which those explanations conform to the covering law
model, which ts biology about as well as it does physics.
Keywords Biological laws Physical laws Covering law Deductive-nomological
Empirical laws A priori laws Dispositions Rosenberg Sober Kitcher
When philosophers of biology attempt to characterize the differences between biology and
physics, they often focus on differences between biological and physical explanations.
These differences are often presented as differences in the degree to which the two kinds of
explanations adhere to the once generally accepted covering law model of scientic
explanation. After all, the story goes, the progenitors and chief early adherents of the
covering law model were all either physicists themselves, or at least much enamored of,
and familiar with, physics. Consequently, it is no wonder that their model of explanation is
a better t for their science of choice than it is for biology. Identifying biological deviations
from this model of explanation, it is said, will therefore also call attention to signicant
differences between biology and physics. However, although there is fairly widespread
consensus that biological explanation is less suited to the covering law model than physical
explanation, there is considerably less consensus about how exactly the two diverge. Much
of this disagreement has focused on the status, or even the purported non-existence, of
biological laws.
J. Press (&)
Philosophy Department, California University of Pennsylvania, 250 University Ave,
California, PA 15419, USA
e-mail: press@cup.edu
1 3
Biol Philos (2009) 24:359374
DOI 10.1007/s10539-007-9096-4
In the rst part of this paper I intend to concentrate on the views of three inuential
philosophers of biology, Alex Rosenberg, Elliot Sober, and Philip Kitcher, along with their
interlocking criticisms of each other. Although these three philosophers hold widely
divergent views on the nature of biological explanations, they share an interest in the
degree of applicability or inapplicability of the covering law model to biology. Rosenberg
sees the least divergence from this model, arguing that biological explanations do appeal to
empirical laws. However, he claims that such laws are comparatively scarce in biology.
Sober claims that, unlike the empirical laws of physics, the laws of biology are a priori.
Kitcher, on the other hand, rejects biological laws altogether.
For the most part, I intend to withhold my own analysis of these positions until the
second part of the paper, in which I will argue that this focus on laws and divergence from
the covering law model has led us astray. Though the covering law model has few true
adherents today, the approach of these philosophers of biology tacitly grants that it applies
reasonably well to physical explanations: well enough, at any rate, for it to make sense to
assert that physical explanations t the model better than biological explanations. In this
paper, I intend to share this assumption. But though I will, as a result, effectively be
defending the covering law models applicability to biological explanations in what fol-
lows, I do not mean to commit myself to that model as anything like a perfect account of
scientic explanation. Rather, the point of defending the applicability of the covering law
model to biological explanation is to argue that, although certain important differences
between physics and biology can indeed be highlighted by differences between physical
and biological explanations, these differences do not come down to differences in the
degree to which those explanations conform to the covering law model. Biological
explanation, I claim, conforms to the covering law model about as well as physical
explanation.
Part I
1.1 A challenge for biological laws
On the face of it, biology does seem short on the sort of laws required by the covering law
model. Alexander Rosenberg has suggested that this is because constant modication of
species by natural selection undermines most attempts to form biological empirical gen-
eralizations. For example, present-day zebras have stripes, in part because they confuse the
visual systems of lions. But Zebras have stripes is not even successful as a ceteris paribus
generalization because, given the constantly changing relationships between zebras, lions,
and their shared environment, this probably has not always been, nor will it probably
forevermore be, the case. A generalization that is currently perfectly serviceable, like
Zebras have stripes, might be confusing or misleading at other times, and is at any rate
almost certainly false, even as a ceteris paribus generalization, when construed as applying
to all zebras existing at all times (Rosenberg 2001a, pp. 739740).
Zebras have stripes might not seem like a particularly good candidate for a biological
law simply because it would be such a low-level law. But the same sort of reasoning will
apply to apparently more robust generalizations. It once would have seemed (had there
been anyone around to whom it might have seemed) a reasonable generalization to say that
reproduction takes the form of mitosis. That would not be a plausible law of biology today,
and before we go formulating some law that would be plausible today, we should consider
what might be the case tomorrow. For, while any attempt to generate laws of biology will
360 J. Press
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be an attempt to cut biological nature at the joints, the problem with biological nature is
that the joints keep shifting places.
Rosenbergs argument has been criticized by Kenneth Waters (1998), on the grounds
that it conates two sorts of biological empirical generalizations. Generalizations like
zebras have stripes describe the distribution of traits among populations and taxonomic
groups, and are indeed constantly modied by natural selection. But the generalizations
involved in, for example, predicting the relative tnesses of horses and zebras in lion-
infested environments neednt quantify over the groups mentioned in these distributional
generalizations. Instead, this second sort of generalization would identify causal regular-
ities between, say, certain types of visual systems and certain types of visual targets,
regardless of how those systems and targets might be taxonomically distributed.
However, in the context of the present question concerning the applicability of the cov-
ering law model to biology, these generalizations identifying causal regularities are no help.
The problem is that the gradual and haphazard nature of natural selection still tends to
produce poorly delineated kinds over which to generalize. Both within and beyond the lion
species lies a near continuumof visual systems, each of which may behave slightly differently
when confronted with zebra stripes. As a result, Waters himself admits that, although these
causal generalizations possess some properties of laws of nature, they fall short of actually
being laws (Waters 1998, p. 29). Consequently, if these generalizations are meant to take the
place of laws in biological explanations, there would still be a difference, albeit perhaps a less
signicant one, in the applicability of the covering law model to physics and to biology.
1.2 Applying the covering law model to biological explanations
Rosenbergs solution to his own problem is to claim that the only laws of biology are the
empirical generalizations that make up Darwins theory of natural selection. It is these
laws, plus the laws of molecular bio-chemistry, upon which biological explanations rely
(Rosenberg 2001b). He axiomatizes natural selection (by his own admission, very roughly)
in the following way:
(1) Biological systems not on the verge of extinction or xity reproduce with heritable
variations.
(2) If heritable variation obtains among biological systems, then there will be tness
differences among the biological systems.
(3) In the long run, the more t variants will leave a higher proportion of descendants
than the less t variants.
(4) [It follows that] until xity or extinction is attained, there will be descent with
modication, i.e. evolution (Rosenberg 2001a, p. 752).
These principles do not fall prey to his criticism of other biological laws since
None of them is subject to qualications or ceteris paribus clauses in virtue of the
operation of selective forces on the earth. After all, these principles constitute the
mechanism of natural selection itself; there is no scope for natural selection to
qualify, limit, or shape its own operation (Rosenberg 2001a, p. 752).
Unfortunately, these laws are not necessarily as broadly applied as Rosenberg claims.
Since (1)(3) are empirical universal generalizations, the argument from (1)(3) to (4) does
conform to the covering law model. It explains a further empirical generalization: that
evolution occurs in all species. However, many biological explanations are more particular
Physical explanations and biological explanations 361
1 3
than this. Biologists explain not only the occurrence of evolution simpliciter, but also the
evolution of this or that trait in this or that population. But, when they use the sort of
reasoning exemplied above for such particular purposes, what goes in the place of (1)(3)
need no longer be universal generalizations. For example,
(1
0
) The monkeys in population P (which is not on the verge of extinction or xity)
reproduce with heritable variations of tail dexterity.
(2
0
) If heritable variation of tail dexterity obtains among the monkeys in P, then the
prehensile-tailed monkeys in P will be tter than the less-dexterous-tailed monkeys in P.
(3
0
) In the long run, the tter, prehensile-tailed monkeys in P will leave a higher
proportion of descendants than the less t, less-dexterous-tailed monkeys in P.
(4
0
) [It follows that] until xity or extinction is attained, the proportion of monkeys in P
with prehensile tails will increase.
In virtue of their reference to a particular population and a particular trait, (1
0
)(3
0
) are
no longer universal, and hence the explanation apparently fails to conform to the covering
law model. And yet, as an explanation, it seems no worse off for this. Of course, it could be
that contemporary evolutionary biologists always come to believe particular propositions
like (1
0
)(3
0
) by deducing them from Rosenbergs laws (1)(3). But, in principle, a pre-
Darwinian (or just very poorly schooled) biologist, unaware of the universal truths (1)(3),
but aware of the particular truths (1
0
)(3
0
), could nevertheless construct this second
explanation. This is because both the inference from (1)(3) to (4) and the inference from
(1
0
)(3
0
) to (4
0
) are underwritten by the same complex model of the process of natural
selection. As it happens, natural selection applies to all living things, but this universal
applicability appears to be irrelevant to explaining its effects in any given case. Conse-
quently, it seems that biological explanations do not necessarily appeal to Rosenbergs
laws of natural selection. At the same time, biological explanations like (1
0
)(3
0
) do not
obviously appeal to the laws of bio-chemistry either, and so it looks as though it fails to
appeal to any of the laws Rosenberg accepts.
Additionally, Philip Kitcher has objected to this sort of axiomatization of natural
selection on the grounds that it fails to accurately describe the scientic advance effected
by Darwin. The problem with taking the main premises of Darwins argument as com-
prising the revolutionary theory of natural selection is that these purported laws are now,
and were in Darwins own time, stunningly obvious. Even biological novices can conrm
that individual members of a species vary, that some of this variation leads to variations in
tness, and that offspring tend to resemble their parents.
But, says Kitcher,
We [legitimately] expect that the fundamental principles of a novel scientic theory
should be those statements introduced by the theory that most stand in need of
defense and conrmation (Kitcher 1985, p. 130).
Basically, Kitchers argument here is that, whereas the introduction of Darwins theory of
natural selection was revolutionary, the laws of natural selection are not at all revolutionary,
and hence the laws of natural selection cannot be the fundamental principles of biology.
1.3 Alternative models of biological explanation
Since the covering law model requires that all explanations involve derivation of the
phenomena to be explained from a set of premises which include empirical laws of nature,
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if the above considerations, or others, decisively rule out the existence of most empirical
biological laws, biological explanation will generally fail to conform to the covering law
model. But if we grant that the explanatory power of physical explanations does come from
something like the covering law model, those who reject that model for biological
explanation need an alternative account of the explanatory power of biology.
1.3.1 A priori laws
One such account suggests that the difference between physical and biological explanation
lies in the fact that, unlike the laws appealed to in the former, the laws appealed to in
biological explanations are a priori. Elliot Sober, for example, has suggested that biological
laws are actually mathematical models. He offers the following as an example of a
mathematical model that might be considered a law of biology.
HardyWeinberg Law: If the frequency of the A gene is p and the frequency of the a
gene is q at some locus in a population, then the frequencies of the three genotypes
AA, Aa, and aa will be p
2
, 2pq, and q
2
, respectively (Sober 2000, p. 73).
This is a law about the frequency of genotypes in a population of randomly sexually repro-
ducing organisms. However, if we strip the law of its biological vocabulary, we are left with:
Mathematical truth: If the frequency of property A is p and the frequency of property
a is q for some set of objects, then (if the possession of A is independent of the
possession of a) the frequencies of the property sets {A, A}, {A, a}, and {a, a}
possessed by those objects will be p
2
, 2pq, and q
2
, respectively.
When the properties are interpreted as genes and the objects are interpreted as organisms,
we get the HardyWeinberg Law. But we could just as well interpret the properties as
Heads and Tails, and the objects as (weighted or unweighted) coin-toss events and then
we would get a law about the frequency of coin-tosses. Either way, what makes the law
true is entirely a priori.
Sober thinks that this just demonstrates an interesting way in which biological laws
differ from physical laws.
In physics, general laws such as Newtons Law of Gravitation and the special theory
of relativity are empirical. In contrast, many of the general laws in evolutionary
biologyseem to be nonempirical (Sober 2000, p. 72).
However, though mathematical models obviously play a signicant role in the biological
sciences, it is not clear that they can assume the explanatory duties of empirical laws. As
Rosenberg argues:
Necessarily true mathematical models will have a role in explanation only to the
degree that they are reected in generalizations that describe actual causal processes
(Rosenberg 2001a, p. 746).
A mathematical model cannot explain or predict the behavior of some biological system
unless we rst know that the model can appropriately model the system. That means we
have to know there is a coherent way of interpreting the various mathematical objects,
properties, and relations that appear in the model as biological objects (species, organisms,
organs etc.), properties (traits, etc.), and relations (descendant/ancestor relations, mating
relations, predator/prey relations, etc).
Physical explanations and biological explanations 363
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Knowing this involves knowing certain empirical facts. If the model is to have general
applicability, then it involves knowing certain general empirical facts. For example, if we
know that, say, all mammal populations satisfy the background conditions of the Hardy
Weinberg Law (i.e. sexual reproduction with random mating), then the HardyWeinberg
Law will allow us to conclude that the genes in mammal populations are always distributed
as it suggests. But if so, it seems that it is this empirical generalization (i.e. that all
mammals satisfy the background conditions) which is the real biological law doing the
explanatory work.
But if it must be the empirical generalizations that serve as laws, we are stuck with
Rosenbergs problem again. There is no guarantee that the assumptions of the Hardy
Weinberg Law will always be satised by mammals, since sexual reproduction may itself
be altered by natural selection or even supplanted by some other form of reproduction. It is
a matter of empirical law whether the model has any general application, and Rosenberg
has claimed that such laws are not in the cards.
On the other hand, just as with Rosenbergs laws of natural selection, one need not seek
general applicability of mathematical models. The HardyWeinberg Law could be applied
just as well to some particular population satisfying the conditions of the model, even if
this satisfaction were a peculiarity or an accident. Something like this seems to be what
Sober has in mind. In a reply to Rosenbergs earlier claim that he had denied the existence
of biological laws altogether, Sober writes
Of course, if one stipulates that laws must be empirical, the [mathematical models]
will not be counted as [laws]I have no strong objection to this usage of the term
law, but the resulting claimthat evolutionary models do not provide laws
gives quite the wrong impression. The point is not that evolutionary biology is
exclusively given over to narratives about historical particulars (Sober 1996, p. 467).
However, if we apply the HardyWeinberg law in this way, nothing explanatory appears to
happen at all. If a biologist knows that, as a matter of, possibly particular, fact, this
population of koala bears reproduces sexually and randomly, it is merely a matter of
calculation, enshrined as the HardyWeinberg Law, that their genes will be distributed as
the law says. It is no more explanatory of the resulting distribution than the fact that I have
two apples in my left hand and three oranges in my right hand, along with the law that
2 + 3 = 5, is explanatory of the fact that I am holding ve pieces of fruit. In either case, the
mathematical law allows us to re-describe a narrative of historical particulars in order to
make explicit certain facts implicit in that narrative, but it does not add to our under-
standing of those particulars, or to the narrative.
1.3.2 Historical biological explanations without laws
Kitchers view of biological explanation arises from his general view of scientic expla-
nation, which conveniently sidesteps the issue of natural laws. On this view (Kitcher 1999)
an argument will form the basis of an explanation when it instantiates an argument pattern
that unies the explanation of many phenomena. In identifying such patterns, we are
supposed to look for patterns of argument that are simultaneously relatively specic and
yet widely applicable to the phenomena to be explained. A theory is a collection of these
patterns that unies the explanations within its domain. Whether or not a science has any
empirical laws becomes a matter of whether universal empirical generalizations make an
appearance in these unifying patterns.
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In evolutionary biology, Kitcher thinks the relevant argument patterns are what he calls
Darwinian histories.
A Darwinian history for a group G of organisms between t
1
and t
2
with respect to a
family of properties F consists of a specication of the frequencies of the properties
belonging to F in each generation between t
1
and t
2
(Kitcher 1985, p. 139).
Unlike the unifying patterns of physics, which frequently incorporate empirical laws,
Darwinian histories make no reference to empirical laws. The general idea is that any
explanation in terms of natural selection will critically involve a recitation of the traits
possessed by the ancestors of the organism, population of organisms, or species possessing
the trait to be explained. Certain kinds of questions, Kitcher says, can be answered by an
appeal to this sort of minimal pattern alone. Other sorts of evolutionary questions might
require that additional detail be added to the pattern. For example, if we want to explain
why gibbons have prehensile tails, we will describe the history of the changing frequency
of the prehensile-tail trait among gibbon ancestors and add a gloss about the tness
advantages conferred upon those ancestors by their prehensile tails.
Much more could be said about the details of Kitchers account. However, the preceding
is enough to show that, elegant as it is, it is insufcient. It is certainly true that scientic
explanations tend to follow patterns, and that explanations that unify a wide range of
phenomena are often highly prized. But the fact that explanations often have the properties
Kitcher identies as relevant does not entail that it is those properties that make them
explanatory. In order to produce an explanation, it is necessary to show that the phe-
nomenon to be explained was, at least to some degree, to be expected. Nothing in Kitchers
Darwinian histories by themselves demonstrates this. As Rosenberg puts it:
If evolutionary explanations really do not appeal implicitly or otherwise to an
ampliative principle of natural selection, the explanatory force of Kitchers
selectionist explanatory pattern remains ungrounded. There is only a narrative
(Rosenberg 2001a, p. 751).
Consider the Darwinian-history explanation of the prehensile tails of gibbons. Many
generations ago, there were more or less gibbon-like organisms that lived in trees. A very
few of these organisms had prehensile tails. Many of these organisms died or were seri-
ously injured as a result of falling from trees, but the organisms with prehensile tails
suffered considerably fewer such incidents than their cousins with less-dexterous tails. In
subsequent generations, the frequency of the prehensile-tail trait increased, and eventually
became more or less universal among the descendants of the rst organisms. Modern
gibbons are descendants of these organisms.
In some sense, this is just the sort of thing one expects from an evolutionary expla-
nation. However, none of these statements is even relevant to the current possession of
prehensile tails by gibbons unless some empirical generalization or other is also true. For
example, in order to understand the relevance of prehensile tails to falling, one must know
that, ceteris paribus, animals with prehensile tails are less likely to fall out of a tree than
those without such tails. In order to understand the relevance of the death and injury
reports, one must know that, ceteris paribus, animals who die or are severely injured tend
to leave fewer offspring than those who do not. And in order to understand the relevance of
all these historical events to modern gibbons, one needs to know that offspring tend to
resemble their parents. The point is that, even if explanations are arguments that t a
pattern, they must surely also be good arguments. Without laws, the explanantia of
Kitchers Darwinian histories fail to entail their explananda.
Physical explanations and biological explanations 365
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Part II
2.1 Re-evaluating differences
The tangle of objections and counter-objections involved in this debate seems to me
to provide sufcient motivation to look more closely at the contention that the best way to
evaluate the differences between biology and physics involves comparing the degree
to which they conform to the covering law model. The alternative accounts of explanation
offered by Sober and Kitcher identify two main possible departures from this model. First,
biological explanations may appeal to a priori laws whereas physical explanations typically
fail to do so, appealing instead to empirical laws (Sober). Second, biological explanations
may fail to appeal to empirical laws, either because they appeal to a priori laws instead
(Sober), or because they do not appeal to laws at all (Kitcher). What I aim to do now is
argue that neither of these purported differences between biology and physics is as deep as
is often supposed.
2.1.1 A priori physical laws
Professor Malcolm Wells, a recently retired physicist at the end of a distinguished career,
still teaches now and then. Truth be told, he has never been entirely satised with his
performance in the classroom, and now that he has the time, he wants to get it right.
The problem has always seemed to be that, no matter how thoroughly he explains
something, he fails to call attention to some crucial bit of background information.
Ruminating on this conundrum, his thoughts drift back to the sixtiesand his under-
graduate philosophy of science course. Philosophical methods, he remembers, have a
remarkable tendency to force one to be explicit. Perhaps if he were to present his lecture on
Snells Law tomorrow as an attempt to ll out a complete covering law explanation,
thorough student comprehension would be ensured.
So, the next day Wells begins by conducting a quick experiment (see Fig. 1). First, he
attaches his laser pointer to the side of a sh aquarium at a xed angle. He turns the pointer
on and marks the point on the bottom of the aquarium where the resultant red dot appears.
He next lls the aquarium with 30 cm of water. As the water rises, the dot moves. After the
tank is lled, he marks the new position of the dot, which has moved 27 cm. Snells Law,
he says, explains this phenomenon. He then writes the following on the board:
(1) Snells Law: If a light ray passes from one medium to another, the light will be
refracted such that the angle of incidence a, the angle of refraction b, and the index of
refraction of the new medium i, are related as sina = isinb.
(2) The laser beam passed from air to water.
(3) The index of refraction for water is 1.333.
(4) The laser pointer is oriented such that a = 60.
(5) The water is 30 cm deep.
(6) Therefore, the red dot moved 27 cm when the water was added.
Condent of having included every essential premise, Wells asks Any questions?
Sadly his condence almost immediately evaporates in a urry of raised hands. Some of
the questions are simply demands for denitions of unfamiliar terms. Wells dutifully
directs the students asking these questions to their textbooks, gently but rmly reminding
366 J. Press
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them of their ability to read. Most of the questions, however, are devoted to calculation.
Some of the students are not sure how to solve the Snells Law equation for b. Others fail
to see how determining the angle of refraction is relevant to the distance the dot traveled.
Though inwardly cursing the public schools lack of emphasis on the practical appli-
cation of mathematics, Wells admits that the students have a point. Covering law
explanations are not generally presented as explicitly including mathematical truths, since
such truths are not supposed to carry any of the explanations content. But that doesnt
mean that one can follow the explanation without knowing the mathematical truths nec-
essary to carry out the derivation. And this calculation isnt completely trivial. So Wells
runs through the calculations, drawing Fig. 1 as he proceeds.
Since we know that a = 60 and that i = 1.333, we can use Snells Law to calculate
that b & 40 by taking the sine of 60, dividing that by 1.333, and taking the cosecant
of the result. Now, suppose we draw a line perpendicular to the surface of the water,
through the point where the laser beam meets the water. Drawing this line constructs two
right triangles. Both triangles have a side lying along the perpendicular and both have a
side lying along the bottom of the aquarium. One triangles hypotenuse is coincident with
the beams path when the aquarium is empty; the others is coincident with the refracted
beams path after the water is added. The difference in the lengths of the bottom sides of
these two triangles will be the distance traversed by the dot. The bottom sides of the
smaller and larger triangles are (30 cm)(tan 40) & 25 cm and (30 cm)(tan 60) & 52
cm, respectively. Consequently, the dot moved 52 cm 25 cm = 27 cm.
But Professor Wells, interjects a student, shouldnt all these facts about calculation
be added to your explanation?
Wells tries for a moment to remember the logical positivists views about empirical vs.
mathematical content, but quickly realizes that, given his present purposes, it doesnt really
matter whether mathematical truths count as a part of the explanation. He is trying to ll in
as many gaps in his students knowledge as possible, so he agrees to put it all up. He adds
several premises to his explanation, including sin 60 & (1.333)(sin 40), (30)(tan
40) & 25, and 52 25 = 27. Condence regained, he again asks, Any questions?
Professor, I understand all the calculations now, but theres still something troubling
me. How do we know that Snells Law is the right equation?
Well, Snells Law tells you how angles of incidence and refraction are related when
light passes from one medium to another, and thats what happened in this case.
40
60
30cm
25cm 27cm
Laser
Pointer
Refracted
Light Ray
Un-
refracted
Light Ray
52cm
Fig. 1 Professor Wells experiment
Physical explanations and biological explanations 367
1 3
How do we know thats what happened?
You watched me do it, unless you were asleep. Its recorded as premise (2).
No sir, I wasnt sleeping, and I see premise (2). Its just that Im not seeing the
relevance of premise (2) to premise (1).
Ahwell(1) makes a hypothetical general claim about what happens any time a
certain sort of event occurs, and (2) says that an event which satises that hypothetical
condition has, in fact, occurred. We can therefore conclude that what the law says
happens in such circumstances will, in fact, happen now. This is just a matter of logic. If
I remember right, the philosophers have a special name for it. Modusponensor
something like that anyway.
Ohwell if it has a special name, I guess it must be OK.
Indeedlets call it a day, shall we?
I take it that the moral of this vignette is relatively obvious. Sober has claimed that at
least one fundamental difference between biological and physical explanations is that only
the former signicantly involve a priori laws. But the story makes clear that, merely in
virtue of being logical derivations, all explanations rely on a priori generalizations. When
explanations are quantitative, many of these generalizations will be mathematical (like the
generalization that whenever one has 52 of something, and then gets rid of 25 of them, one
is left with 27). But even purely qualitative explanations must at least rely on the logical
truths that link their premises (like modus ponens). So, far from identifying a unique
feature of biological explanation, Sober has identied a feature that is omnipresent in
explanation, especially if explanation is to be conceived on the covering law model, or on
any model that represents explanations as arguments.
At the same time, Sober is onto something. While both physical and biological
explanations appeal to universal a priori generalizations, such generalizations often seem
to assume greater importance in biology. Consider another of Sobers examples. R. A.
Fisher explained why populations of sexually reproducing organisms tend to have male/
female ratios close to 1:1. Suppose that there is a signicant deviation from a 1:1 sex
ratio in a population, and that there is heritable variation in the sex ratios of the offspring
individual organisms produce. In such a situation, individuals who produce more off-
spring of the minority sex will have more grandoffspring than individuals who produce
more offspring of the majority sex (Each individual in the grandoffspring generation
requires a pairing of male with female, so offspring of the minority sex will necessarily
participate in more pairings than offspring of the majority sex). Consequently, whenever
the sex ratio of the population deviates from 1:1, there will be a selection bias in favor of
production of the minority sex until such time as the balance is redressed (see Sober
2000, p. 17).
This explanation appeals to various empirical considerations. I would even contend that
it appeals to empirical laws. I intend to say more about what these laws might be in the
next section, but for now it will be sufcient to note that identication of these empirical
considerations is not Fishers main contribution to biology here. It is relatively easy to
conrm that many organisms reproduce sexually, that their pairings are often more or less
random, that the sex ratio of their offspring is heritable and varies, and so on. Instead, the
challenging part is realizing that organisms which t this description are overwhelmingly
likely, as a matter of mathematical necessity, to give birth to an even balance of males and
females. If we try to t Fishers explanation into the covering law model (using as yet
unspecied empirical laws), Fishers main contribution seems to be recognizing the
inference pattern, rather than recognizing some new empirical fact or law.
368 J. Press
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This is signicantly different from the Snells Law explanation. Professor Wells expects
that listing the empirical laws and empirical particular facts necessary to deduce the
movement of the dot will provide a sufcient explanation, and he seems within his rights to
be a bit vexed when his students fail to make the necessary mathematical and logical
connections. Surely a good physics student, given the empirical laws and facts, could at
least manage to work through the trigonometry and algebra on his or her own time (The
student who fails to grasp modus ponens may seem positively surreal unless one has
attempted to teach introductory logic). It is virtually impossible to imagine an alternate
scientic history in which the community of physicists comes to realize the truth of Snells
Law without yet realizing its applicability to something like Professor Wells experiment.
But something analogous appears to happen in biology all the time. I conclude that Sober is
more or less right to focus on the role of a priori generalizations in characterizing the
difference between physics and biology, but wrong to think that the difference is a matter
of substituting a priori laws for empirical ones. Both physical and biological explanations
depend upon the truth of both empirical and a priori generalizations, but, unlike physical
explanations, the Eureka! premise of biological explanations tends to be a priori.
I think this difference also accounts for Kitchers reluctance to accept that Darwins
great achievement was just the introduction of the laws of natural selection. These
empirical generalizations were old news, hence Kitcher claims they could not have rev-
olutionized biology. However, as Kitcher himself says, What was in dispute in the
Darwinian revolution was not so much the truth of [the laws of natural selection], as their
signicance, (Kitcher 1985, pp. 130131). Darwins achievement was in demonstrating
that the familiar process of natural selection was sufcient, by itself, to produce the entire
biological world. Darwin was lling in an explanatory gap analogous to the one Professor
Wells attempts to ll by explicitly listing mathematical truths. Just as most of Wells
students are capable of doing their own trigonometry, most of Darwins contemporaries
understood the logic of natural selection. But, just as many of Wells students failed to see
how to apply their knowledge of trigonometry, Darwins contemporaries often failed to see
natural selection when and where it was at work. In physics, lling in the inferential details
may tend to be grunt work (although this is certainly not always true). But what we have
seen is that lling in the inferential details in biology often leads to signicant discoveries.
Ironically, it is Kitchers insistence that the fundamental principles of a novel scientic
theory should be those statements introduced by the theory that most stand in need of
defense and conrmation, (Kitcher 1985, p. 130), which seems to be an artifact of the
early covering law theorists preoccupation with physics. The inferences of natural
selection, being a priori, may not have been in need of defense and conrmation, but it
did take the genius of Darwin to demonstrate their importance.
Thus, I conclude that Sobers recognition of the importance of a priori mathematical
models in biology does not necessarily undermine the applicability of the covering law
model to biology. Physical explanations, just as much as biological explanations, rely upon
the truth of a priori generalizations, but in neither case do these generalizations ll the role
of empirical laws.
2.1.2 Empirical biological laws
However, this similarity between physical and biological explanations wont save the equal
applicability of the covering law model to both sorts of explanation unless they both also
rely upon empirical laws. So far, the only empirical biological laws that seem plausible are
Physical explanations and biological explanations 369
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Rosenbergs laws of natural selection. Though the considerations above appear to deect
Kitchers historical objection to those laws, we have already seen that explanations of
particular evolutionary events, as opposed to the general fact of evolution, do not appear to
essentially rely upon those laws. Combined with Rosenbergs concerns about the shifting
joints of biological kinds, this might lead us to doubt whether there are enough biological
laws to satisfy the demands of the covering law model.
Consider again the explanation of prehensile tails. There do not appear to be any
obvious laws hiding amongst its premises. The explanation contains assertions that in this
particular population the prehensile-tail trait is heritable, that there is variation with respect
to possession of this trait, and that those with the trait are tter than those without it. Since
these claims are all claims about a particular population and a particular trait, they lack the
generality of laws. If this explanation contains empirical biological laws, they are hiding.
One possibility is that the explanation relies on implicitly assumed, well known
laws. Uncovering such implicit laws is a common method of deecting apparent counte-
rexamples to the covering law model (Hempel 1966, pp. 5152). However, in the current
case, no such lurking premises appear to be necessary. If these particular facts about
particular monkeys and their tails are true, it is a matter of mathematical necessity that the
frequency of the prehensile-tail trait will probably increase in the population.
So, if there are empirical generalizations at work in this explanation, they must be
hiding in the explicitly stated premises. Consider the claim that the prehensile-tail trait is
heritable. This is a claim that the members of this population have a certain disposition (or
propensity): the disposition of tending to pass the prehensile-tail trait to their offspring.
Explicitly dening the concept of a disposition is a notoriously vexing task (for example,
see Malzkorn 2001) that I dont intend to undertake here. However, for present purposes I
think we can make do with the relatively safe assertion that claims about complex, higher-
level dispositions (i.e. dispositions like heritability or tness, perhaps unlike fundamental
dispositions such as electrical repulsion or proton decay) imply the claim that the dispo-
sition occurs because of some set of underlying natural regularities. We would not call
some objects tendency to behave in a certain way a disposition if we believed that this
tendency were a mere coincidence or miracle. These implied natural regularities are
empirical laws if anything is (Of course, some philosophers might want to reject talk of
laws altogether, but since what is at issue in the present instance is whether or not bio-
logical explanations conform less well to the covering law model than physical
explanations, and since any outright rejection of natural laws would equally undermine the
covering law model for both sorts of explanation, I will assume that talk of laws is
acceptable for present purposes).
One interesting property of these disposition claims is that, although they appear to have
this implication, one can intelligibly make a disposition claim without necessarily being
able to specify which laws underwrite the disposition. For example, Kepler, Newton, and
Einstein all equally shared the belief that planets are disposed to travel in elliptical paths.
But whereas Newton thought this disposition a result of a natural law that universally
attributes an attractive force to massive objects, Einstein thought it underwritten by a law
governing the effects of massive objects on the curvature of space-time. Kepler lacked any
understanding of the underlying regularities at all, but certainly believed that they were
there. His belief counts as a belief in the disposition of planets to orbit in ellipses merely in
virtue of the fact that he believed there to be some set of underlying laws responsible for it.
The same was importantly true of heritability. Darwin famously lacked any theoretical
understanding of why organisms tend to produce offspring that resemble them. But he
realized that due to some sort of underlying natural process, organisms do have this
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tendency. Luckily, this was all he needed to know about heritability to construct natural
selection explanations. In the explanation of prehensile tails, so long as the prehensile-tail
trait is heritable, it doesnt much seem to matter why it is heritable. All that is important is
that heritability is a natural disposition, rather than a coincidence.
Assertions of tness are also disposition claims. A claim that one type of organism is
tter than another implies a claim that, certain of the laws of nature being what they are,
individuals of the former type are likely to leave more offspring than individuals of the
latter type. Once again, one might believe that one type of organism is tter than another
without necessarily having an understanding of precisely which laws of nature are
responsible.
I think this is where our missing empirical laws have been hiding. Though it is true
enough that we can explain the evolution of prehensile tails entirely in terms of the
properties possessed by particular organisms, some of these properties are dispositions.
These disposition claims imply that there are certain unspecied universal natural laws
which underwrite the dispositions. Clearly, if known, these unspecied laws would allow
us to construct covering law explanations of the dispositions themselves, and these
explanations could then be substituted into whatever higher-level explanations refer to
those dispositions. In the case of heritability, for example, we now know enough about the
underlying genetic processes to at least sketch a (bio-chemical) covering law explanation.
But the great value of dispositional language is that it allows us to refer to these laws
without specifying them, or even knowing precisely what they are. The attribution of
dispositions thus acts as a sort of placeholder for the unspecied laws in a covering law
explanation by placing outer bounds on which natural laws might be true. The explanation
of prehensile tails depends on the laws of nature being the sort of laws that, in the
conditions that obtain, give rise to heritability of that trait in gibbon ancestors. Since
heritability is easily veriable in this case, whatever the laws of nature in fact are, they
must be of the right sort. Similar limitations are placed on the laws by other disposition
claims in the explanation, such as attributions of tness. Since the explanation relies on
these laws without specifying them, we can explain why it looks as though the biologists
explanation makes no reference to laws at all, even though it in fact conforms to the
covering law model.
This defense of the covering law model involves a greater degree, as it were, of
implicitness than Hempels. Whereas his defense claims that covering law explanations
can leave certain known premises about laws implicit, I am suggesting that the attribution
of dispositions allows us to implicitly include in our explanations laws about which we are
ignorant or which we understand only incompletely. After all, the point of including laws
in covering law explanations is that this ts the phenomenon to be explained into a pattern
of uniformities and shows that its occurrence was to be expected, (Hempel 1966, p. 50).
So long as our knowledge of dispositions constrains the ways that the underlying laws
might be sufciently that, however they actually are, the phenomenon to be explained was
to be expected, this purpose will have been fullled.
The same defense can be made, I suspect, of any true biological explanation. Fishers
sex ratio explanation, for example, contains references to dispositions (both heritability and
tness), so it implicitly makes reference to unspecied laws. Furthermore, biology is rife
with attributions of other dispositions. Hearts are disposed to pump blood, wolves are
disposed to kill deer, and so on. Wherever explanations rely upon claims about disposi-
tions, they rely upon laws, and this satises the covering law model.
This also allows us to avoid Rosenbergs worry about the historical contingency of
biological kinds. Rosenberg argues that empirical generalizations cannot be laws because
Physical explanations and biological explanations 371
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they appeal to historically dened categories that allow for signicant temporal variation in
traits. But as we have seen, evolutionary biologists dont really need to appeal to gener-
alities concerning all members of a species. For example, if we want to explain why zebras
are harder for lions to catch than horses, we will be able to do so by pointing out that
present-day zebras have stripes, present-day horses dont, and present-day lions have
trouble seeing stripes. Over time, zebras, horses, and lions may change in ways that
undermine these generalizations, but that doesnt matter since these are not the laws to
which the explanation appeals. Instead, the explanation appeals to the laws, whatever they
may be, that govern the assorted complicated processes which underlie the disposition of
the visual systems of lions to detect horses more effectively than zebras. These laws do not
have to be explicitly mentioned in the explanation, or even known by the biologist who
develops it, so long as it is clear that the visual systems of lions do, in fact, possess this
disposition. The biologists assertion that this disposition exists places bounds on the laws
that might exist, and this is enough to satisfy the covering law model.
This is consistent with Rosenbergs claim that whenever biological explanations fail to
refer to the laws of natural selection, they refer to the laws of molecular bio-chemistry, and
that biology is thus reducible to bio-chemistry (Rosenberg 2001b). However, my thesis is
only tangential to the issue of reductionism. First, while it may be that disposition claims in
biological explanations always act as placeholders for bio-chemical laws, if it turned out
otherwise, reductionism might be in trouble. But my thesis about the equal applicability of
the covering law model to both biology and physics would not be similarly imperiled, since
it makes no assertions about the sorts of laws underlying the disposition claims in bio-
logical explanations. Second, if, as Rosenberg maintains, the issue of reducibility comes
down whether or not the explanations of biology can be complete without including
reference to bio-chemical laws, my own view suggests that this issue is subject to an
ambiguity. For, while I claim that biological explanations, via their disposition claims,
refer to laws of nature, I do not require that these laws be made explicit, or even that they
be known, so long as the disposition claim sufciently constrains the possible ways the
laws might be. So, if by complete, we mean that an explanation is fully explicit, then I
would agree that biological explanations cannot be complete without a full statement of the
laws upon which they rely. But we might simply mean by complete that an explanans
contains all that is needed to deduce its explanandum. In this sense, I think that biological
explanations that leave all reference to the laws of nature implicit in disposition claims can
still be complete, since their disposition claims imply that the laws, whatever they may
actually be, must be such that the explanandum was to be expected.
2.1.3 The covering law model and approximation
Some may doubt, however, that these references to unknown laws hidden in disposition
claims really satisfy the covering law models demand for covering laws. Leaving known
laws implicit (ala Hempel) is one thing, but if disposition claims merely imply that there
are some underwriting laws or other, at best one has provided only the promise of an
explanation.
But the covering law model has always been intended as an idealization. Besides
explanations with implicit laws, Hempel also allows deviations from the ideal that he calls
partial explanations (Hempel 1962, pp. 1618). For example, he considers a case in
which Freud explained a slip of the pen as the result of wishful thinking. In this case,
although the strongest law endorsed by Freud would be a law to the effect that strong
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unconscious desires will cause a slip that symbolically fullls those desires in some way or
other, Freuds explanation is of a particular slip (a misdated letter). The explanation is
partial in the sense that, while it succeeds in predicting a slip, it fails to predict the
particular sort of slip that actually occurred.
Something similar happens when explanations involve quantitative approximations. If
Professor Wells initial measurements involve some margin of error, that margin will be
transmitted to the explanandum. Just as Freuds explanation entailed only that some sort of
slip would occur, Wells explanation from approximated initial conditions entails only that
the laser dot will move some distance falling in a small range around 27 cm. So this also
seems to count as a partial explanation in Hempels sense, and, while we might on that
account want to say that it fails to be an ideal explanation, it would certainly seem overly
fastidious, even within the covering law tradition, to deny that Wells had explained the
motion of the dot.
The biological explanations that I claim invoke unknown laws via disposition claims do
not merely involve this sort of approximation of initial conditions. However, I believe we
can say that they involve an analogous approximation of laws. If biological disposition
claims implied nothing but the existence of some natural law or other, then biologists really
would be out of the explanation business. But as I have said above, the attribution of a
disposition also places a constraint on what the underwriting laws must be like: the laws
must be laws that will actually bring the disposition about. For example, the disposition of
monkeys to be more successful in reproduction when they manage to avoid falling out
of trees places certain constraints on the law of gravity. That monkeys have this dispo-
sition does not tell us that objects in freefall at the Earths surface have an acceleration of
9.8 m/s
2
, but it does tell us that the acceleration of gravity at the Earths surface is
sufcient to accelerate a monkey who falls from the height of a tree to a velocity high
enough to kill or seriously damage it when it nearly instantaneously decelerates upon
reaching the ground. The existence of the same tness disposition also places limits on the
laws governing the forces that account for the strength and exibility of monkey bones, the
effectiveness of various tissue repair mechanisms, and much more.
These constraints may often be fairly weak, but the point is that they appear to be strong
enough to produce (partial) biological covering law explanations. Just as the explanation of
the motion of the laser dot can survive some imprecision in its prediction resulting from
approximated initial conditions, so the explanation of prehensile tails can survive
approximated laws. When made fully explicit, the disposition in the prehensile tail
explanation would be cashed in, not for claims about the precise set of laws from which
prehensile tails may be deduced, but instead a range of sets of laws, any one of which
predicts the evolution of prehensile tails. Since any one of the sets of laws leads to the
same prediction, we can show that, even given what little knowledge of laws is contained
in our knowledge of the dispositions, we should expect prehensile tails. Of course, just as
with other sorts of partial covering law explanations, we could, and sometimes do, produce
a better explanation by more precisely pinning down what has previously only been
approximated. As we improve our knowledge of physics, chemistry, and molecular biology
we can narrow down the range of covering laws to which our explanation appeals. But we
neednt necessarily do so to have a partial covering law explanation just as good in its own
way as Professor Wells explanation of the laser dot or Freuds explanation of his slip of
the pen.
So once again we nd that the differences between biological and physical explanations
are merely a matter of degree. All sciences employ approximations of one sort or another
in the construction of covering law explanations. In virtue of these approximations, the
Physical explanations and biological explanations 373
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explanations in which they occur fall short of perfect realization of the covering law model,
but still seem worthy to be called (partial) explanations. As one might expect, biologists,
who deal with extremely complex systems, will need to rely relatively heavily on various
sorts of approximation if they are to explain anything at all. This will include approxi-
mation of the covering laws through attributions of dispositions. But in so doing, they are
not falling signicantly shorter of the covering law ideal than scientists in other disciplines,
and they need not be construed as engaged in some wholly different sort of explanatory
activity.
Conclusion
Sober and Kitcher have indeed uncovered important differences between physical and
biological explanations. Scientic progress in biology appears to rely more heavily on
recognition and application of a priori principles or models governing explanatory infer-
ences, and less heavily on discovery of new empirical laws or particular facts, than does
scientic progress in physics. However, these differences should not be characterized as
differences in the degree to which biological and physical explanations conform to the
covering law model of scientic explanation. For, while a priori principles do appear to
play a larger role in biological explanations than in physical explanations, such principles
are essential components of any covering law explanation, including physical covering law
explanations. And, while biological explanations do appear to place less emphasis on
empirical laws, and often omit explicit mention of them, reference to empirical laws still
plays an essential role in such explanations, just as it does in physical explanations. The
appearance that biological explanations make no reference to empirical laws is the result of
the fact that the laws involved in biological explanations are typically only implied by
premises attributing dispositions, which place bounds upon the laws of nature without
actually asserting the truth of particular laws.
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