Thursday, December 28, 2017
The cog-neuro of plants
There are many (e.g. Erich Jarvis) who think that the basic
hierarchical properties of language are direct reflections of its vocalic
expression. This is what makes ASL and other signed languages so useful. The
fact that they exist and that modulo their manual articulation they look, so
far as we can tell, just like any other language (see here
for discussion) puts paid (and paid in full!) to any simple minded idea that
linguistic structure is “just” a reflection of their oral articulation.
Why do I mention this? Because I have just been reading some
popular pieces about a potentially analogous case in neuroscience. Let me
explain.
Several years ago I read a piece on plant “neurobiology” in
the New Yorker penned by Michael
Pollan (MP) (here).
The source of the quotes around ‘neurobiology’ is a central concern of the
article in that it explores whether or not it is appropriateto allow that plants may have a cognitive
life (‘bullshit’ is one of the terms tossed around) or whether this is just
another case of metaphors run amok. One large influential group of critics
thought the idea obscurantist bordering on the incoherent (see here).
Here’s a quote from the MP piece quoting one of the critics Lincoln Taiz:
Taiz says that the writings of
plant neurobiologists suffer from “over-interpretation of the data, teleology,
anthropomorphizing, philosophizing and wild speculations. (6)
Wow, this is really bad, as you can tell when the charge is “philosophizing”!!!
Can’t have any of that. At any rate, the main reason for Taiz’s conclusion, it
appears, is that plants do not have brains (an uncontroversial point) or
neurons (ditto). And if one further assumes that brains with neurons are
required for cognition of any kind then the idea that plants might have memory,
might use representations, learn and make context sensitive judgments is simply
a category mistake. Hence the heat in the above quote.
In a recent Aeon essay,
Laura Ruggles (LR) reprises the issues surrounding thinking plants (here).
It appears that things are still contentious. This does not really surprise me.
After all, the idea that plants cognize really is a weird and wonderful
suggestion. So, that it could be false or, at the least, ill supported, strikes
me as very plausible. However, as the quote above indicates, this is not the
nature of the criticism. The objection is not that the evidence is weak but
that the very idea is incoherent. It is not false. It is BS. It is not even
high class BS, but a simple category mistake due to bad anthropomorphic
philosophical speculation generated by teleologically addled minds. Needless to
say, I got interested.
Why the heat? Because it directly challenges the reductive
neurocentric conception of cognition that animates most of contemporary
cog-neuro (just as ASL challenges the vocalic conception of grammar). And it
does so in two ways: (i) It reflects the strong commitment of the methodology
of “neuron doctrine” in cog-neuro and (ii) It reflects the strong commitment to
the idea that biological memory and computation supervenes on a connectionist
architecture (i.e. the relevant computations are inter-neural rather than
intra-neural). Let me say a word or two about each point.
The “neuron doctrine” is the idea “that cognitive activity
can be accounted for exclusively by basic neuroscience. Neuronal structure and
function, as identified by neurophysioplogy, neuroanatomy and neurochemistry,
furnish us with all we need to appraise the animal mind/brain complex” (see here: 208).
This idea should sound familiar because it is the same one that we discussed in
the previous post (here).
The position endorses reductionist methodology to the study of the brain (more
accurately, a methodological monism), one that sees no fruitful contribution
from the mental sciences to cog-neuro. KGG-MMP rehearses the arguments against
this rather silly view, but it clearly has staying power. Marr fought it in the
1980s and his proposal that problems in cog-neuro need to be tackled at (at
least) three different levels (computational, algorithmic/representational,
implementational), which interact but are distinct and provide different kinds
of explanatory power is a response to precisely this view. This view, to repeat,
was prevalent in his day (well over 30 years ago!) and is still going strong,
as witnessed by the fact that KGG-MMP felt the need to reiterate his basic
arguments yet again.
The plant “neuroscience” debate is a manifestation of the
same methodological dogmatism, the position that takes it for granted that we
once we understand neurons, we will understand thought. But it is even more of
a challenge to this neurocentric view. If plants can be said to cognize (have
representations, memories, process information, learn) then not only is the
methodological thesis inappropriate, but the idea that cognition reduces to
(exclusively lives on) neuronal
structure, is wrong as well (again, think ASL and vocalization wrt grammar). If the plant people are onto something
then the having memories is independent of having brains, as is learning and representation
and who knows what else. So if the plant people are right, then not only is the
neuron doctrine bad methodology, it is also ontologically inadequate.
None of this should be surprising if you have any
functionalist sympathies. As Marr noted, the materials that make up a chess
board/pieces can vary arbitrarily (wood, marble, bread, papier mache) and the
game remains the same (the rules/game of chess is readically independent of the
physical make-up of the board and pieces). Whether the relation of cognition to
brains is more like the chess case or less is an open question. One view
(Searle comes to mind) is that no brains, no cognition. On this view, the connection
between brain structure and cognition is particularly tight in that the former
is a necessary feature of the latter (thinking can only live in brains (though, IMO, there is more than a touch of
mystical vitalism in Searle’s position)). If the plant cognitivists are right,
then this is simply incorrect.[1]
In sum, though metaphysical reduction does not lend credence
to methodological reduction, if even the former is untenable, then it is quite
implausible that the former can stand. Why import neuronal methodological dicta
in the study of cognition if cognitive machinery need not live in neuronal
wetware?
The neuron doctrine has a more specific expression in todays
cog-neuro. It’s the claim that the brain is basically a complex neural net and
that memory, learning, cognition are products of such neural nets. In other
words, the prevalent view in contemporary cog-neuro is that cognition is an
inter-neuronal phenomenon not an intra-neuronal one. Brains are the locus of
cognition because it brains have inter-neuronal connections. Memories, for
example, are expressed in connection weights and learning amounts to adjusting
these inter-neuronal weights. The plant stuff challenges this view as well. How
so? Because if plants to cognize they seem to do it without anything analogous to a brain (more exactly, this
assumption is common ground in the discussion). So, if plants have memories
then it looks like they encode these within
cells. LR mentions epigenetic memory as a possible memory substrate. These
involve “chromatin marks,” which “are proteins and small chemical groups that
attach to DNA within cells and influence gene activity” (LR: 3). This mechanism
within cells suffices to physically implement “memory.” And if this is so, then
it would provide evidence for the Gallistel-King conjecture that memories can
be stored biochemically within cells. Or to state this more carefully: if
plants can code memories in this way, why not us too and maybe neuronal
connectionism is just a wrong-headed assumption, as Gallistel has been arguing
for a while. Here is MP making this point:
How plants do without a
brain…raises questions about how our brains do what they do. When I asked
Mancuso about the function and location of memory in plants, he…reminded me
that mystery still surrounds where and how our memories are stored: “it could
be the same kind of machinery, and figuring it out in plants may help us figure
it out in humans.” (MP:19)
Indeed.
Ok, there is lots of fun stuff in these essays. It is fun to
see how plant people go about arguing for mental capacities in plants. There
are nice discussions of experiments that appear to show that plants can
“habituate” to stimuli (they pretend-drop plants and see how they react), can
“learn” new associations (use wind as conditioned stimulus for light) among
stimuli, and can to anticipate what will happen (where sun will be tomorrow) in
the absence of the thing being anticipated (in the absence of input from the
sun), which suggest that plants can represent the trajectory of the sun. Is
this “true” and do plants cognize? I have no idea. But an a priori denial that
it is possible is based on conceptions of what proper cog-neuro is that we have
every reason to reject.
[1]
So too if machines can cognize (Searle’s target), something that seems less
challenging for some reason than that plants do. There is some nice speculation
in the MP article as to why this might be the case.
Wednesday, December 20, 2017
More on modern university life
Universities are spending more and more money on administrative staff. Here is a post with references to more in depth material that puts some numbers to this process. Administration is eating up all the revenue and it is growing faster than any other part of the university. Three points of interest in the post: first, faculty positions have risen in line with student numbers (56% rise in students and 51% rise in faculty). The out of proportion rise lies with administrators and their staffs. It has exploded. Second, this trend is bigger in private universities than public ones. The post notes that this "looks to be the opposite of what we would expect if it were public mandates lying behind this [i.e. rise in bureaucrats, NH] trend. Third, this really is a new trend. Universities are changing. As The post notes:
...in the "good old days" top admins tended to be more senior faculty with reasonably distinguished records who had been on campus for a long time and knew the people and the place. Now we have undistinguished professional managers...I don't know about where you are, but this seems to pretty well sunup the state of play at those institutions that I am acquainted with (like my own).
Monday, December 11, 2017
How to study brains and minds
There is currently a fight going on in cog-neuro whose
outcome GGers should care about. It is illuminatingly discussed in a recent
paper by Krakauer, Ghazanfar, Gomez-Marin, MacIver and Poeppel (KGG-MMP) (here).
The fight is about how to investigate the mind/brain connection. There are two
positions. One, which I will call the “Wrong View” (WV) just to have a useful
mnemonic, takes a thoroughly reductionist approach to the problem. The idea is
that a full understanding of brain function will follow from a detailed
understanding of “their component parts and molecular machinery” (480). The
contrary view, which I dub the “Right View” (RV) (again, just to have a name),[1]
thinks that reductionism will not get nearly as far as we need to go and that
the only way to get a full understanding of how brains contribute to
thinking/feeling/etc. requires neural implementations in tandem with (and more
likely subsequent to) “careful theoretical and experimental decomposition of
behavior.” More specifically, “the detailed analysis of tasks and of the
behavior they elicit is best suited for discovering component processes and
their underlying algorithms. In most cases,…the study of the neural
implementation of behavior is best investigated after such behavioral work” (480). In other words, WV and RV differ
not over the end game (an understanding of how the brain subvenes the brain
mechanisms relevant to behavior) but the best route to that end. WV thinks that
if you take care of the neuronal pennies, the cognitive dollars will take care
of themselves. The RV thinks that doing so will inevitably miss the cognitive forest
for the neural trees and might in fact even obscure the function of the neural
trees in the cognitive forest. (God I love to mix metaphors!!). Of course, RV
is right and WV is wrong. I would like to review some of the points KGG-MMP
makes arguing this. However, take a look for yourself. The paper is very
accessible and worth thinking about more carefully.
Here are some points that I found illuminating (along with
some points of picky disagreement (or, how I would have put things differently)).
First, framing the issue as one of “reductionism” confuses
matters. The issue is less reduction than it is a neurocentric myopia. The
problem KGG-MMP identifies revolves around the narrow methods standard practice
deploys not the ultimate metaphysics that it endorses. In other words, even if
there is, ontologically speaking, nothing more than “neurons” and their
interactions,[2]
discovering what these interactions are and how they combine to yield the
observed mental life will require well developed theories of this mental life expressed
in mentalistic non-neural terms. The problem then with standard practice is not
its reduction but its methodological myopia. And KGG-MMP recognizes this. The
paper ends with an appeal for a more “pluralistic” neuroscience, not an
anti-reductionist one.
Second, KGG-MMP gives a nice sketch of how WV has become so
prevalent. It provides a couple of reasons. First, has been the tremendous
success of “technique driven neuroscience” (481). There can be no doubt that
there has been an impressive improvement in the technology available to study
the brain at the neuronal level. New and better machines, new and better
computing systems, new and better maps of where things are happening. Put these
all together and it is almost irresistible to grab for the low hanging fruit
that such techniques bring into focus. Nor, indeed should this urge be
resisted. What needs resisting is the conclusion that because these sorts of
data can be productively gathered and analyzed that these data suffice to
answer the fundamental questions.
KGG-MMP traces the problem to a dictum of Monod’s: “what is
true of the bacterium is true of the elephant.” KGG-MMP claims that this has
been understood within cog-neuro as claiming that “what is true for the circuit
is true for the behavior” and thus that “molecular biology and its techniques
should serve as the model of understanding in neuroscience” (481).
This really is a pretty poor form of argument. It
effectively denies the possibility of emergence. Here’s Martin Reese (here)
making the obvious point:
Macroscopic systems that
contain huge numbers of particles manifest ‘emergent’ properties that are best
understood in terms of new, irreducible concepts appropriate to the level of
the system. Valency, gastrulation (when cells begin to differentiate in
embryonic development), imprinting, and natural selection are all examples.
Even a phenomenon as unmysterious as the flow of water in pipes or rivers is
better understood in terms of viscosity and turbulence, rather than
atom-by-atom interactions. Specialists in fluid mechanics don’t care that water
is made up of H2O molecules; they can understand how waves break
and what makes a stream turn choppy only because they envisage liquid as a
continuum.
Single molecules of H2O do not flow.
If one is interested in fluid mechanics then understanding will come only by
going beyond the level of the single molecule or atom. Similary if one is
interested in the brain mechanisms underlying cognition or behavior then it is
very likely that we will need to know a lot about how groups of fundamental
neural elements interact, not just how one does what it does. So just as a
single bird doesn’t flock, nor a single water molecule flow, nor a single
gastric cell digest, so neither does a single brain particle (e.g. neuron)
think. We will need more.
Before I get to what more, I should add here that
I don’t actually think that Mondo meant what KGG-MMP take him to have
meant. What Monod meant was that the principles of biology that one finds in
the bacterium are the same as those that we find in the elephant. There is
little reason to suppose, he suggested, that what makes elephants different
from bacteria lies in their smallest parts respecting different physical laws.
It’s not as if we expect the biochemistry to change. What KGG-MMP and Reese
observe is that this does not mean that all is explained by just understanding
how the fundamental parts work. This is correct, even if Monod’s claim is also correct.
Let me put this another way: what we want are explanations.
And explanations of macro phenomena (e.g. flight, cognition) seldom reduce to
properties of the basic parts. We can completely understand how these work
without having the slightest insight into why the macro system has the features
it does. Here is Reese again on reduction in physics:
So reductionism is true in a
sense [roughly Monod’ sense, NH]. But it’s seldom true in a useful sense.
Only about 1 per cent of scientists are particle physicists or cosmologists.
The other 99 per cent work on ‘higher’ levels of the hierarchy. They’re held up
by the complexity of their subject, not by any deficiencies in our
understanding of subnuclear physics.
So, even given the utility of understanding the
brain at the molecular level (and nobody denies that this is useful), we need
more than WV allows for. We need a way of mapping two different levels of
description onto one another. In other words, we need to solve what Embick and
Poeppel have called the “granularity mismatch problem” (see here ).
And for this we need to find a way of matching up behavioral descriptions with
neural ones. And this requires “fine grained” behavioral theories that limn
mental mechanisms (“component parts and sub-routinges”) as finely as neural
accounts describe brain mechanisms. Sadly, as KGG-MMP notes, behavioral
investigation “has increasingly been marginalized or at best postponed”
(481-2), and this has made moving beyond the WV difficult. Rectifying this
requires treating behavior “as a foundational phenomenon in its own right”
(482).[3]
Here is one more quibble before going forward. I am
not really fond of the term ‘behavioral.’ What we want is a way of matching up
cognitive mechanisms with neural ones. We are not really interested in
explaining actual behavior but in explaining the causal springs and mechanisms
that produce behavior. Focusing on behavior leads to competence/performance
confusions that are always best avoided. That said, the term seems embedded in
the cog-neuro literature (no doubt a legacy of psychology’s earlier
disreputable behaviorist past) and cannot be easily dislodged. What KGG-MMP
intends is that we should look for mental
models and use these to explore neural models that realize these mental
systems. Of course, we assume that mental systems yield behaviors in specific
circumstances, but like all good scientific theories, the goal is to expose the
mental causes behind the specific behavior and it is these mental causal
factors whose brain realization we are interested in understanding.The examples KGG-MMP gives show that this is
the intended point.
Third, KGG-MMP nicely isolates why neuroscience needs mental
models. Or as KGG-MMP puts is: “Why is it the case that explanations of
experiments at the neural level are dependent on higher level vocabulary and
concepts?” Because “this dependency is intrinsic to the very concept of a
“mechanism”.” The crucial observation is that “the components of a mechanism do
different things than the mechanism organized as a whole” (485). As Marr noted,
feathers are part of the bird flight mechanism, but feathers don’t fly. To
understand how birds fly requires more than a careful description of their feathers.
So too with neurons.
Put another way, as mental life (and so behavior) is an
emergent property of neurons how
neurons subvene mental processes will not be readily apparent by only studying
neural properties singularly or collectively.
Fourth, KGG-MMP gives several nice concrete examples of
fruitful interactions between mental and neural accounts. I do not review them
here save to say that sound localization in barn owls makes its usual grand
appearance. However, KGG-MMP provides several other examples as well and it is
always useful to have a bunch of these available on hand.
Last, KGG-MMP got me thinking about how GGish work
intersects with the neuro concerns the paper raises, in particular minimalism
and its potential impact for neuroscience. I have suggested elsewhere (e.g. here)
that MP finally offers a way of bridging the granularity gap that Embick and
Poeppel. The problem as they saw it, was that the primitives GGers were
comfortable with (binding, movement, c-command) did not map well to primitives
neuro types were comfortable with. If, as KGG-MMP suggests, we take the notion
of the “circuit” as the key bridging notion, the problem with GG was that it
did not identify anything simple enough to be a plausible correlate to a neural
circuit. Another way of saying this is that theories like GB (though very
useful) did not “dissect [linguistic, NH] behavior into its component parts or
subroutines” (481). It did not carve linguistic capacity at its joints. What
minimalism offers is a way of breaking GB parts down into simpler
subcomponents. Reducing macro GB properties to products of simple operations
likeMerge or Agree or Check Feature
promises to provide mental parts simple enough to be neurally interpretable. As
KGG-MMP makes clear finding the right behavioral/mental models matters and
breaking complex mental phenomena down into its simpler parts will be part of
finding the most useful models for neural realization.
Ok, that’s it. The paper is accessible and readable and
useful. Take a look.
[1]
As we all know, the meaning of the name is just what it denotes so there is no
semantic contribution that ‘wrong’ and ‘right’ make to WV and RV above.
[2]
The quotes are to signal the possibility that Gallistel is right that much
neuronal/cognitive computation takes place sub
neuronally.
[3]
Again, IMO, though I agree with the thrust of this position, it is very badly
put. It is not behavior that is foundational but mentalistic accounts of
behavior, the mechanisms that underlie it, that should be treated as
foundational. In all cases, what we are interested in are the basic mechanisms
not their products. The latter are interesting exactly to the degree that they
illuminate the basic etiology.
Friday, December 1, 2017
Fodor and Piatelli Palmarini on Natural Selection
The NYT obit on Jerry Fodor accurately recognizes the important contributions he made to philosophy, psychology and linguistics. The one reservation noted, the strained reception of his late work on evolution and his critique of Darwin. It accurately notes that Jerry saw the achilles heal of natural selection theories residing in their parallels with behaviorism (a parallel, it should be noted, that Skinner himself emphasized). Jerry and Massimo concluded that to the degree the parallels with behaviorism were accurate then this was a problem for theories of natural selection (a point also made by Chomsky obliquely in his review of Skinner at the outset of the cognitive revolution). I think it is fair to say that Jerry and Massimo were hammered for this argument by all and sundry. It's is one thing to go after Skinner, quite another to aim to decapitate Darwin (though how much Darwin was a radical selectionist (the real target of Jerry's and Massimo's critique) is quite debatable). At any rate, as a personal tribute to the great man I would like to post here an outline of what I took to be the Jerry/Massimo argument. As I note at the end, it strikes me as pretty powerful, though my aim is not to defend it but to elucidate it for most of the critiques it suffered did not really engage with their claims (an indication, I suspect, that people were less interested in the argument than in defending against the conclusion).
The content of the post that follows was first published in roughly this form in Biolinguistics. I put it up here for obvious reasons. Jerry Fodor was a great philosopher. I knew him personally but not as well as many of my friends did. I was charmed the few times I socially interacted with him. He was so full of life, so iconoclastic, so funny and so generous (most of his insights he graciously attributed to his grandmother!). I looked up how often I talked about Jerry's stuff on FoL and re-reading these made me appreciate how much my own thinking largely followed his (though less funny and less incisive). So, he will be missed.
So, without further ado, here is a reprise of what I take to have been the Jerry/Massimo argument against Natural Selection accounts of evolution.
***
Jerry Fodor and Massimo Piatelli-Palmarini (F&P, 2010) have recently
argued (in What Darwin Got Wrong) that
the theory of Natural Selection (NS) fails to explain how evolution
occurs. Their argument is not with the
fact of evolution but with the common claim that NS provides a causal mechanism
for this fact. Their claim has been
greeted with considerable skepticism, if not outright hostility.[1] Despite the rhetorical heat of much of the
discussion, I do not believe that critics have generally engaged the argument
that F&P have actually presented. It
is clear that the validity of F&P’s argument is of interest to
biolinguists. Indeed, there has been
much discussion concerning the evolution of the Faculty of Language and what
this implies for the structure of Universal Grammar. To facilitate evaluation of F&P’s
proposal, the following attempts to sketch a reconstruction of their argument
that, to my knowledge, has not been considered.
1. 'Select' is not 'select for', the latter being intensional.[2]
2. The free rider problem shows that NS per se does not have
the theoretical resources to distinguish between ‘select’ and ‘select for.’
3. If not, then how can NS causally explain evolutionary change?
4. There are two ways of circumventing the free rider problem.[3]
a.Attribute mental powers to NS, i.e. NS as Mother Nature,
thereby endowing NS with intentionality and so the wherewithal to distinguish
‘select’ from ‘select for.’
b.Find within NS Law supporting counterfactuals, i.e.
Laws of Natural Selection/Evolution, which also would suffice to provide the
requisite intentionality.
5. The first option is clearly nuts, so NS accounts must be presupposing
4b.
6. But NS contains no laws of evolution, a fact that seems to be widely
recognized!
7. So NS can't do what it purports to do; give a causal theory that
explains the facts of evolution.
8. Importantly, NS fails not
because causal accounts cannot be given for individual
cases of evolution. They can be and routinely are. Rather the accounts are
individual causal scenarios, natural histories specific to the case at hand,
and there is nothing in common across the mechanisms invoked by these individual
accounts besides the fact that they end with winners and losers. This is, in
fact, often acknowledged. The only relevant question then is whether NS
might contain laws of NS/Evolution? F&P argue that NS does not
contain within itself such laws and that given the main lines of the theory, it
is very unlikely that any could be developed.
9. Interestingly, this gap/(flaw) in NS is now often remarked in the Biology
Literature. F&P review sample some work of this sort in the book. The
research they review tends to have a common form in that it explores a variety
of structural constraints that were they operative would circumscribe the
possible choices NS faces. However, importantly, the mechanisms proposed are
exogenous to NS; they can be added to it but do not follow from it.
10. If these kinds of proposals succeed then they could be combined with
NS to provide a causal theory of evolution. However, this would require giving
up the claim that NS explains evolution.
Rather, at most, NS + Structural
Theories together explain evolutionary change.[4]
11. But, were such accounts to develop the explanatory weight of the
combined 'NS + Structural Theory' account would be carried by the added structural
constraints not NS. In other words, all that is missing from NS is that part
that can give it causal heft and though this could be added to NS, NS itself
does not contain the resources to develop such a theory on its own. Critics might then conclude as follows: this
means that NS can give causal accounts when supplemented in the ways
indicated. However, this is quite
tendentious. It is like saying Newton's
theory suffices to account for electro-magnetic effects for after all Newton's
laws can be added to Maxwell's to give an account of EM phenomena!
12. F&P make one additional point of interest to linguists. Their review and conclusions concerning NS
are not really surprising for NS replays the history of empiricist psychology,
though strictly speaking, the latter was less nutty than NS for empiricists had
a way of distinguishing intentional from non-intentional as minds are just the
sorts of things that are inherently intentional. In other words, though attributing mental
intentional powers to NS (i.e. Mother Nature) is silly, attributing such powers
to humans is not.
This is the argument. To be honest, it strikes me as pretty
powerful if correct and it does indeed look very similar to early debates
between rationalist and empiricist approaches to cognition. However, my present intention has not been to
defend the argument, but to lay it out given that much of the criticism against
the F&P book seems to have misconstrued what they were saying.
[1] See,
for example: A misguided attack on
evolution, Massimo Pigliucci. 2010. Nature 464, A misunderstanding Darwin, Ned Block and Philip Kitcher. 2010. Boston
Review of Books, 35(2), Futuyma, D. 2010, Two
critics without a clue. Science, 328: 692-93.
[2]
Intensional contexts are ones in which extensionally identical expressions are
not freely interchangeable. Thus, if
John hopes to kiss Mary and Mary is The Queen of the Night, we cannot conclude
that John hopes to kiss the Queen of the Night.
[3]
F&PP develop this argument in Chapter 6.
The classic locus of the problem is S.J. Gould and R.C. Lewontin. The
spandrels of San Marco and the Panglossian paradigm: a critique of the
adaptationist program. Proceedings of
the Royal Society of London, Series B biological sciences, vol 205, 1979,
581-98.
[4] Observe, the supposition that selection is simply
a function of “external” environmental factors lies behind the standard claim
that NS (and NS alone) explains why evolutionary changes are generally
adaptive. Adding structural “internal”
constraints to the selective mix, weakens the force of this explanation. To the degree that the internal structural
factors constrain the domain of selection, to that degree the classical explanation
for the adaptive fit between organism and environment fails.
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