Please Note: This article was previously published under the title, “Whole Organisms and Their Evolutionary Intentions — An Overview”.
Every organism is continually dying in order to live. Breaking-down activities are prerequisites for building up. Complex molecules are synthesized, only to be degraded later, with their constituents recycled or excreted. In multicellular organisms such as vertebrates, many cells must die so that others may divide, differentiate, and proliferate. Many cancers reflect a failure to counterbalance proliferation with properly directed death processes.
You and I have distinct fingers and toes thanks to massive cell death during development. The early embryo’s paddle-like hands give way to the more mature form as cells die and the spaces between our digits are “hollowed out”. In general, our various organs are sculpted through cell death as well as cell growth and multiplication. During development the body produces far more neurons than the adult will possess, and an estimated ninety-five percent of the cell population of the thymus gland dies off by the time the mature gland is formed.1
Despite all this life and death, I doubt whether anyone would be tempted to describe the embryo’s cells as “red in tooth and claw”. Nor do I think anyone would appeal to “survival of the fittest” or natural selection as a fundamental principle governing what goes on during normal development. The life and death of cells appears to be governed, rather, by the developing form of the whole in which they participate.
But this has been a truth hard for biologists to assimilate, since it has no explanation in the usual causal sense. One way to register the problem is to ask yourself what you would think if I suggested that organisms in populations thrive or die off in a manner governed by the evolutionary outcome toward which they are headed — that the pattern of thriving and dying off becomes what it is, in some sense, because of that outcome. It is not a thought any evolutionist is likely to tolerate. But perhaps the occasional intrepid researcher will be moved to inquire: “Why not?” After all, we can also ask about the cells populating our bodies: do they thrive or die off in a manner governed by the forthcoming adult form? And here the answer appears to be a self-evident “yes”.
Perhaps, when we have come to accept what we see so clearly in individual development, we will find ourselves asking the “impossible” question about evolutionary development: Does natural selection really drive evolution, or is it rather that the evolving form of a species or population drives what we think of as natural selection? Are some members of an evolving species — just as with the cells of an embryo’s hands — bearers of the future, while other members, no longer being “fit” for the developing form of the species, die out?
What makes this idea seem outrageous is the requirement that inheritances, matings, interactions with predators, and various other factors in a population should somehow be coordinated and constrained along a coherent path of change. Unthinkable? But the problem remains: Why — when we see a no less dramatic, life-and-death, future-oriented coordination and constraint occurring within the populations of cells in your and my developing bodies — do we not regard our own development as equally unthinkable?
Once we are willing to consider them, “unthinkable” ideas may sometimes reveal profound truths. My thesis will be that when we look at how organisms realize their distinctive forms and ways of life — when we look at all the features of biological activity in general, without ignoring the inconvenient ones — we can recognize, in the facts we already have, that evolution has a living, well-organized, well-coordinated, well-directed character analogous to that of individual development. The conclusion isn’t even speculative. It requires no new or unexpected discoveries. It is simply what we find ourselves looking at, if we have not turned our eyes away.
This is not to say that the direction of evolution is owing either to an external guiding power, such as a breeder, or to a conscious “aiming” or planning. Rather, the evolutionary narrative arises from the agency and developmental powers of organisms and communities of organisms, as they express their own character and realize their potentials in the presence of the prevailing environmental challenges and opportunities.
Before we get into it, however, it would be good to have in our minds a picture drawn from life. I have chosen a 1927 description by the British naturalist and ornithologist, Edward Max Nicholson.2
A scene from life
“The male must leave the flock, if he has belonged to one, and establish himself in a territory which may at the time be incapable of sustaining him alone, but must later in the season supply a satisfactory food-supply for himself, his mate and family, and for as many birds of other species as overlap his sphere of influence. He must then sing loudly and incessantly for several months, since, however soon he secures a mate, trespassers must be warned off the territory, or, if they ignore his warning, driven out.
“His mate must help with the defence of the territory when she is needed; pairing must be accomplished; a suitable site must be found for the nest; materials must be collected and put together securely enough to hold five bulky young birds; eggs must be laid in the nest and continuously brooded for a fortnight till they hatch, often in very adverse weather; the young are at first so delicate that they have to be brooded and encouraged to sleep a great part of the time, yet they must have their own weight of food in a day, and in proportion as the need of brooding them decreases, their appetites grow, until in the end the parents are feeding four or five helpless birds equal to themselves in size and appetite but incapable of digesting nearly such a wide diet.
“Enemies must be watched for and the nest defended and kept clean. When the young scatter, often before they can fly properly, they need even greater vigilance, but within a few days of the fledging of the first brood a second nest will (in many cases) be ready and the process in full swing over again. All this has to be done in face of great practical difficulties by two creatures, with little strength and not much intelligence, both of whom may have been hatched only the season before.”
That phrase, “and not much intelligence”, is perhaps ill-advised. It may be that in some sense we can say that this bird, typically weighing less than one ounce (18 – 29 grams), doesn’t possess a great deal of intelligence. But, at the same time, who will deny the stunning wisdom playing through all the multi-faceted performances and accomplishments — the physiology, development, behavior, and entire life story — of this feathery little creature? Perhaps we need to distinguish between the intelligence an organism possesses, and that by which it is possessed — between the intelligence it consciously exercises (if any), and that which runs deeper than our own reflective awareness. Or again: between the intelligence employing a brain, and the still more profound intelligence capable of forming that brain.
Lungs develop in the human embryo in order to breathe air that is not yet there, just as eyes develop before there is light to see. Birds build nests as a preparation for egg laying, and then raise chicks toward maturity. Monarch butterflies migrate thousands of miles southward from Canada in order to overwinter in Mexico. Amoebas move toward food particles as a preparation for engulfing and digesting them. Future requirements and possibilities somehow play into the present as tasks are accomplished, efforts rewarded, needs fulfilled, and interests pursued, from feeding and mating to metabolism and DNA replication.
We know that in all living activity earlier performances are coordinated in such a way as to prepare for later ones. And the continual adjustments in response to disturbances along the way tell us that this intricate coordination is not accidental, but rather belongs to a consistent pattern through which the meaning of the activity comes to successful expression. That is, the consistency of pattern lies, not in any necessity implied at the outset by prearranged physical relationships, but in the significant ends or purposes being realized. Nothing in the the detailed physical conditions of a fertilized egg cell — or of any later stage of growth — dictates that a given liver cell at time A must divide and differentiate so as to produce a particular new kind of cell that should migrate to position X in response to a lesion inflicted by a parasite. It happens, rather, because of the need for healing and the natural movement toward wholeness.
Material resources produced by an organism’s activity in the past may provide both necessary and constraining conditions under which it will work in the future. But the living activity itself is not driven from the past in this way. It is purposive — or, more broadly, is a working out of the organism’s way of being, of the meanings at play in its life. And therefore the activity always has something of an end-directed or future-oriented character. The elaboration of meanings in time, like any story, always holds together as a forward movement.
Every child can grasp it: the meaning of living activity gives narrative significance to — and, at times, in a rough sort of way, may predict — a sequence of events. It is what makes sense (including scientific sense) of whatever is happening. That’s why we always think of our pets as doing something, even if what they are doing is resting. There is good evidence that all biologists “get” this intuitively. But the sense-making, teleological aspect of biological activity is not thought to have a respectable place alongside other modes of scientific explanation.
There are, however, no obstacles to characterizing this central aspect of biology once we are willing to look at it. In 1929 the British physiologist and psychologist, William McDougall, who had earlier succeeded to William James’ chair at Harvard, summarized certain typical features of purposive behavior.3 He was writing specifically about human behavior, but we can recognize these features in all biological activity, conscious or otherwise:
• Goal-directed activity is adaptable to one degree or another. If one strategy fails, the organism may vary it or switch to a different strategy.
• Goal-directed activity tends to be persistent and may be repeatedly renewed even after being effectively blocked for a time.
• As soon as the goal is reached, that particular goal-directed activity ceases.
We do not find the same combination of features in the inanimate world. Yet anyone with a pet dog or cat takes them for granted, with paradigmatic examples being the quest for food and the processes of development.4 If we cannot make sense of something an animal is doing, our first question is likely to be, “Why is it doing that?” — with what purpose? or toward what end? — which is not something we ask of rocks and waterfalls. In our interactions with other creatures we encounter one meaningful life story after another, through which we come to recognize the specific ways of being, the needs, the interests — the meaningful worlds of experience — that all this adaptable and persistent, future-oriented coordination is giving expression to.
Molecules under discipline. Something similar becomes evident even at the molecular level, as the distinguished, twentieth-century cell biologist and National Medal of Science recipient, Paul Weiss, showed us. He spoke of “micro-indeterminacy” and “macro-determinacy”. That is, in the “heaving and churning” cell little if anything is predictable according to a machine-like logic. “The detailed states and pathways of the components not only are so erratic as to defy definition, but, even if a Laplacean spirit5 could trace them, would prove to be so unique and nonrecurrent that they would be devoid of scientific interest”6 — devoid of interest, that is, for anyone who wants the microscopic details to control or explain the meaningful behavior of the whole.
In other words, the cell as a whole exhibits its own specific character and regularity despite the virtually infinite degrees of freedom collectively possessed by its molecular constituents within their watery milieu. The detailed activity, with all its variability and lack of rigid constraint, turns out to be disciplined in a well-directed way toward fulfilling the needs of the cell. The details somehow participate in and reflect their larger context, although not because they are driven to that participation by physical necessity.
And the same principle holds when we shift levels and look at whole cells in relation to still higher states of organization. For example, if the experimentalist removes a limb bud from an embryonic amphibian, mixes up the entire cluster of cells, and then restores the now disordered group of cells to its proper context in the embryo, a normal limb will still develop. Extreme positional freedom among those cells is compatible with the ultimately reliable formation of the limb as a whole. In organisms more generally, an astonishing degree of cell-to-cell variation gives way to remarkably coherent results at the level of tissues and organs.
Weiss summarizes the matter this way:
A unit retains its unity by virtue of the power of subordination which it exerts upon its constituent elements in such a manner that their individual activities, instead of being free and unrelated, will be restrained and directed toward a combined unitary resultant.7
This is as much as to say: the character of a living context shapes and informs its parts, and cannot be understood as merely a deterministic result of those parts. In their own right, the laws governing the parts simply are not capable of expressing, moment by moment, the meaning of the activity in which the parts are caught up.
And so, while no one talks in a natural way about animal behavior without alluding to its meaning and purpose, the more impressive fact may be that no one can avoid talking in the same way about living activity at the molecular level. “Impressive”, I say, because this was the level at which the ultimate reality of “blind mechanism” was supposed to be established. Yet, despite that expectation — and seemingly without even noticing the irony — molecular biologists have routinely defined their subdisciplines and research projects in the only way possible: by referring to the meaning and end-directed character of the activity they are investigating. How does the cell accomplish the task of DNA replication, or the repair of DNA lesions? How does a cell divide? How does it produce proteins? How does it derive energy for its actions through metabolism? And how does it regulate all these activities in relation to the needs of the whole cell and organism?
Here’s just one example. A current challenge embraced by molecular biologists is to understand how hundreds of diverse and diffusible molecules in a watery medium come together and coordinate their interactions in order to carry out the intricate, extended narrative of RNA splicing. In this process they must remove sections of a complex RNA molecule and “stitch” the remaining pieces together in the extremely precise manner required to obtain a functional result. It must all be accomplished in just the right way to yield (through additional, equally elaborate processes) the exact form of the specific protein required right now, in this cell, as opposed to the somewhat different form that may be required later or in a different cell.
There you see Weiss’ principles of micro-indeterminacy and macro-determinacy on vivid display. If we had to explain RNA splicing merely by summing up the individual, law-like behaviors of those hundreds of molecules, with all their degrees of freedom, we would know beyond any doubt: the exponentially multiplying random molecular deviations from the elaborate and drawn-out task at hand would quickly reduce the entire process to a chaotic mess so far as that task was concerned. This is simple physics and chemistry, which were not “made” to sustain meaningful narratives.
No activity is living except by virtue of its task-oriented, end-directed, context-dependent, and meaningful character. Without this, we have physics and chemistry, but not biology. And when we face up to this defining nature of living activity, we can no longer entertain an evolutionary theory that ignores the organism’s agency — an agency whose whole nature is to transform its present existence directionally toward the fulfillment of future potentials.
Agency is not mechanism. In citing the organizing, or coordinating, power evident in all biology, I have just now referred to the organism’s agency. Alternatively, I could have spoken of the intentional (or narrative) context of its life. The idea is merely to provide, one way or another, a name for what we hope to understand more fully. But meanwhile, we have no reason to ignore what we do know — what all our observation shows us — when we turn to other topics such as evolution.
Surely we should offer some sort of name — choose your own, if you wish — for whatever accounts for the organizing of reliable developmental and behavioral narratives. To leave something unnamed is to exclude it from science. And the coordinated, end-directed activity being excluded in this case is not only evident from observation; and it not only poses what seems a significant puzzle (which ought to interest every scientifically minded observer); it also appears on its face to be definitive of living things.
We know enough, however, to reject some explanations for the organism’s agency. There can be no doubt — despite a certain common abuse of language — that organisms are not machines, and they are not executing some kind of program. Unlike machines, organisms grow and continually transform and even heal their own parts. It is hard to see how such parts could “mechanistically” explain their own creation.
The functioning of a machine results from the way the assembly of its preexisting parts was intentionally coordinated by an outside agent in the past. By contrast, the performance of an organism is at every moment an internal coordinating activity through which the ever-changing and maturing parts must be continuously integrated and re-integrated into the developing whole. If the organism were a machine, it would be a different machine at every point in its development — different not merely in the way a computer program’s activity differs from moment to moment, but also different (to use a remarkably inapt expression) in its “hardware”, just as a butterfly differs from its worm-like larva.
Moreover, at the sub-cellular level we see molecules moving and interacting within a fluid medium in order to carry out carefully sequenced narratives — tasks so complex that they challenge our most sophisticated abilities to unravel and articulate their endless nuances. These narrative achievements, which might seem to require a remarkable and practiced synchronization of activities, are accomplished, as we saw a moment ago, despite the fact that the innumerable molecules involved possess many degrees of freedom as they diffuse through the cell’s plasm. And also despite the fact that the context-sensitive task at hand is never exactly the same in any two of the trillions of cells in our bodies, or at any two moments of a several-decades-long life. The rigidly defined and consistent structural constraints necessary for rendering the programmed operation of a computer reliable and mechanistic are altogether absent.
The “mechanistic” causes to which biologists so often appeal are caught up in the highly coordinated activity of an organism’s development. They do not explain their own coordination or their end-directedness. This truth, which we will try to keep in view as we move along, places a huge question mark alongside just about the entirety of current evolutionary theory. We can hardly help doubting a theory that claims to explain changes in organisms over time while ignoring the fundamental fact that biological processes in the present are always in some sense — and in a meaningful (“thoughtful”), non-machine-like manner — orienting themselves toward a not-yet-realized future.
(For a much more expansive assessment of the machine-model of organisms, see “Biology’s Shameful Refusal to Disown the Machine-Organism”, and also the various articles listed under the heading, machine-idea in the topical index for the Biology Worthy of Life project. And for an evaluation of the common argument that evolution somehow explains the organism’s purposiveness, see “Evolution and the Purposes of Life”.)
No one will dispute that a wolf’s development, proceeding from a fertilized egg cell through embryonic and fetal stages to the pup’s birth, and then on through maturation to adulthood, is highly directional. Yes, it’s a path full of unpredictable variation, never exactly repeated in different wolves. But this makes it all the more impressive that the entire trajectory remains persistently wolf-like despite all the adjustments to disturbances and despite all the adaptations to changing conditions. The individual wolf, embedded within its physical and social environment, exhibits its own sort of organizing power, and is capable of negotiating its own, wolf-like path through the exigencies of life.
The ten-day-old heart of the embryonic wolf differs dramatically from the six-week-old heart, which in turn differs from the heart immediately following birth, and this again differs from the heart of the mature wolf. Every biologist will expect the transformations from one stage to another to show all the features of organic activity. The development, persistent and adaptive, will proceed as an intentional narrative, with earlier conditions responded to in such a way as to ensure a workable path toward later achievements. It is just a fact (as I tried to illustrate in the preceding section) that all living activity has this future-oriented aspect.
I doubt whether anyone would want to suggest that there are ways to get from the ten-day-old heart to the mature heart via any pathway not intentionally narrated in the sense of all development.
But suppose we look at an evolutionary sequence, such as the classic textbook lineage of the horse. How might we imagine that a heart, structured that way fifty million years ago in the dog-sized horse ancestor, Eohippus, becomes this heart, structured this way in the Triple Crown winner, American Pharoah? Since every difference in structure must be reflected in how the one organ relates to the rest of the animal — and since all the other parts are also evolving — the question is how this highly integrated, whole-organism transformation could possibly take place.
Can we realistically picture these mutually adjusting metamorphoses being achieved by processes fundamentally less seamless and integral, or less consistent with the general character of all living activity, than the developmental transformations bridging the differences between, say, a two-month- and five-month-old horse embryo?
It is, after all, the whole nature of a developmental narrative to proceed from here to there — to wisely improvise now in the face of unpredictable circumstances, doing so in just the way required for reaching a future state. We would need a powerful and unexpected set of arguments to show that nature, employing any conceivable set of historical processes, could effectively transform such a developmental narrative other than by fully entering into the natural terms of it. We should never forget (although a great deal of evolutionary theory encourages the forgetting): the only biological transformation that ever contributes to evolutionary change is that which occurs within the lives and developmental life cycles of specific organisms.
It is true that evolution, by definition, concerns possibilities of change that go beyond the relatively well-defined stages of any particular organism’s development. And it is true, as we will see later, that evolution has a much broader and more collective intentional focus than we find in individual development. But, as we will also see, this does not allow us to assume an explanatory stance that ignores the nature of organic activity.
The lessons of development. Consider the relatively few cell types in a very young human embryo. These proceed along pathways of differentiation leading eventually to the hundreds of distinct cell types in our mature bodies, from retinal cells to those of heart muscle, from liver to bone, from brain to pancreas. The differences between these cell types are, in their own way, as great as the differences between any two kinds of organism.
Yet cellular transformations do not in general arise from gene mutations. They reflect, among other things, the extraordinarily diverse ways an animal can employ the single genome it was born with.8 And neither do the transformations result from competition for survival between cells of differing fitness. As noted in the introduction, what happens is governed, rather, by the developing form of the organism as a whole.
Evolution: Three Classic Premises
By refusing to acknowledge the living and organizing capacities of organisms, biologists were left with a problem: how to contrive a theory of evolutionary transformation in the absence of any transforming agency. The theory they came up with required a special — one could almost say “magical” — understanding of genes and fitness, each figuring in a foundational premise:
• Genocentrism. Controlling and shaping genes were the decisively important bearers of heredity, the innate causes of traits, and also the substrate for mutational change and evolutionary variation. At the heart of evolutionary theory, they served as proxies for organisms and their living activity. And so the organism’s agency as a self-transforming power was, in effect, reassigned to a single type of molecule, DNA — which, as various biologists have periodically pointed out, is, by itself, one of the most inert molecules in our bodies.9
• Fitness. Some organisms, by virtue of the fitness of their inherited genes, were judged to be more capable than others of surviving and reproducing in their prevailing environment. The resulting differential survival of organisms over time altered the distribution of genes (and gene-combinations) within a population. And because genes stood as proxies for organisms, the evolution of organisms came to be defined by many as “the changing distribution of genes in a population”. This was the key to natural selection, or “survival of the fittest”, as articulated within the Modern Synthesis that brought together Darwin and Mendelian genetics. The principle of selection, then — despite making no reference to any power of coherent transformation in the organism — became the primary agent through which new kinds of organism evolved from previous kinds.
These two premises brought along with them a third:
• Stability. Beneficial, heritable mutations, if they are to contribute to evolution, must be capable of passing down through many generations unaltered. If they were not stable — if they were too quickly lost due to further change — then the processes by which patterns of life and death might spread selected mutations through a population, raising that population’s overall fitness, would not have time to play out. The gene, whether mutated or otherwise, is, we have been told, the overwhelmingly most important thing we know possessing the required stability.
The picture during development, then — and keep in mind that all evolutionary change must be realized in development — suggests truths starkly at odds with the first two premises mentioned in Box 1 at right. That is, it leads us to wonder why anyone would insist on the necessary role of either gene mutations or fitness differences in producing organic change worthy of an evolving species. (On the idea of fitness, see the section, “Fitness differences are not the key to significant evolutionary change”, below.)
Further, the remaining premise — the necessity for stable mutations in order for evolution to occur — also begins to look doubtful. The metamorphosis and diversification of cells teaches us that stability of inherited features is a peculiar criterion for organic change. In the differentiating cell lineage of a developing organism, we see one cell generation succeeding another, not by preserving crucial change, but by compounding it — changing again what has just been changed.
In other words, many of these changes are not heritable over any large number of cellular generations — and had better not be, precisely because the cell lineage is “on the way to somewhere”, proceeding directionally along a pathway of change. This shows how differently we must regard evolutionary processes as soon as we are willing to acknowledge the organism’s agency. The only transformative biological activity we ever witness is thoroughly caught up in this agency.
Yet, as a result of their powerful theoretical bias against the idea of agency, theorists thinking about inheritance and evolution have long avoided taking up the tools necessary to investigate whole-cell inheritance, as opposed to genetic inheritance.10 That is, they have not aimed to discover, in an evolutionary context, anything like the processes whereby change in differentiating cells is compounded upon change — or, more generally, the processes whereby organisms, living in time, routinely and in all their activity, are always transforming themselves toward the future.
The whole organism and its inheritance. Given that many animal bodies consist of hundreds of distinct cell types as different from one another as a liver cell and a retinal cell, any talk about the phenotype of such animals seems a bit odd. This in turn suggests that overly rigid references to the genotype are peculiar as well. It is obvious enough that the organism makes a very different thing of the “same” cellular DNA in different cells. What is most striking here is not some fixed genetic resource, but rather the organism’s evident power of creative organization and coordination. Through this power – and not through any creative capacity inherent in the bits of stuff we call “genes” — the infinitely complex flows of activity in each cell shape themselves, and are shaped, to a particular, specialized outcome in each type of cell.
And what other than this same power of the living organism as a whole coordinates all those unique cells, integrating them into the effective unity of an American Pharoah? Still further, we can ask: How many features of the adult horse are already present in its single-celled origin? Above the molecular level, virtually none. Where, then, do we find the power not only to sustain the organic unity of the whole at any particular moment, but also to bring about the global transformations through which one kind of wholeness passes successfully, from day to day, week to week, and year to year, into a different sort of wholeness, from the earliest embryonic stages to the later adult stages, and all the way to senescence?
Anyone tempted to scorn references to the “power of a living whole” might want to pause a moment and reflect upon the directions in which contemporary biologists are being driven. It is no accident that references to the “context-dependence” of specific molecular interactions have now become almost clichés. Or that the focus of molecular biological research has been shifting inexorably from individual “controlling” causes to flexible and robust networks that in turn are somehow made to fulfill the specific requirements of their larger context. Or that “systems biology”, for all its current mechanistic distortions, has become a major discipline in its own right.
What is really meant by “the system”, or “the context”, if such terms do not signal a nascent (if still largely repressed) recognition that the usual bottom-up, causal style of explanation doesn’t yet bring us to an understanding of distinctively biological narratives?
These more recent currents within mainstream biology may have failed so far to overcome the longstanding habit of ignoring the living being in favor of strictly physical and chemical processes. But they do indicate the direction in which the biology of the future seems destined to move.
The picture we have, then, is one of an organism that can create radically different phenotypes within its own body. These cellular phenotypes are directionally achieved along differentiating cell lineages — and, at a more complex level, we can say something similar about tissue and organ phenotypes. Further, all these divergent types are stably and integrally bound together into the coherent life of one particular creature. And, finally, this creature as a whole proceeds through continual transformation from the earliest embryo onward — all while managing to preserve the unique qualitative substance and character of its kind as it persists and adapts through all the vicissitudes of its existence.
This description belongs to what it means to be a living thing. We are looking at clearly directed activity whose fundamental character, without which it would not be living, enables it to embody a creative relation between past, present, and future, even as it declares itself a master generator of diverse phenotypes, modified as necessary in response to environmental conditions. That all this creative functioning of the organism — exactly the sort of functioning that, on the face of it, seems to bear directly on evolution — should nevertheless be declared more or less irrelevant to the problem of species transformation looks stunningly wrong-headed.
Of all the cellular phenotypes, it would be hard to find one whose differentiation and specialization is more distinctive or more expertly and intricately contrived than the germ cells of sexually reproducing organisms. We can hardly help acknowledging that parental organisms, in carrying out meiosis, genetic recombination, and mating, play a massive role, not only in preserving the genome, but also in transforming it. Deeply embedded in time and always facing the future, every sexually reproducing animal expresses its future orientation most immediately and vividly in the gametes whose full “self-realization” belongs to the next generation.
The entire drama of the germline has been rapidly revealing itself in recent years as a remarkable focus of the organism’s creative “attention”. Are we to believe, then, that this is the one cell lineage in which the organism’s normal, future-oriented activity goes silent? Or that, with all the organism’s expertise at producing, adapting, and stably maintaining diverse phenotypes even without changes in DNA sequence, it “refuses” to employ this expertise when it comes to the preparation of inheritances? Or that the power with which the organism conforms all its cells, tissues, and organs to a unified and integral whole adapted as far as possible to current conditions is a power lost to it in the management of its own germline?
We need to look. No one should think that contemporary biologists have actually spent time looking for anything resembling directed, whole-cell and whole-organism inheritance. Even to suggest the need for it is likely to produce a charge of obscurantism: “How”, the question comes, “can we look at wholes, taking in everything at once?” Yes, we may indeed complain of the fact that life is so integral and complicated — so life-like. But, as I have just pointed out, the need to look at functional wholes is exactly what biologists are already having to face up to as they try to understand cell differentiation and development. This effort certainly requires a great deal of analysis, but the analysis and the integral vision are not the same thing.
When evolutionists finally decide that the “impossible holistic task” is worth pursuing — namely, to observe heritable differences as differences in whole-cell organization and performance, much as developmental biologists today are being forced to learn about the differences in differentiating cell types — then (if they can also avoid the misguided fixation upon endlessly stable variation) they will find themselves possessed of a basis for understanding evolution that bears little relation to the mathematized, gene-centric approach that has ruled for so long.
Actually, the evidence that the answers lie in such a holistic task are everywhere, not just in the experience of developmental biologists with cells. Evolutionary theory has been broadening in every possible direction. To mention a few buzzwords and phrases (no need for the non-specialist reader to worry about these), we hear prominently of niche construction; developmental constraints; transgenerational epigenetic inheritance; lateral gene transfer; endosymbiosis; the microbiome; the organism’s active management of transposable elements (“jumping genes”); the role of culture, learning, and unconscious experience; and, most generally, phenotypic plasticity as a factor capable of leading rather than following natural selection.
What remains missing, however, is what counts most: an explicit acknowledgment of the wise and purposive agency everyone at least unconsciously recognizes in the organism. Without this, we are still left with organisms that are nothing but playthings of imagined causal mechanisms, whether those mechanisms happen to impinge upon their lives from inside or outside their skin. In this sense, even the newer developments (important as they may be for the future) have so far left us with the “same old evolutionary biology”, which is a refusal of the organism’s life.
None of this is to deny real distinctions between individual and evolutionary development. In the latter case we see (in those organisms reproducing sexually) a continual merging of separate lineages. There is also the fact of hybridization across species, genera, and even families. And some of those features just mentioned above — symbiosis of various sorts, cultural inheritance, niche construction, and lateral gene transfer via viruses and microorganisms — also serve to remind us that, while the communal context can be vitally important even for individual development, it becomes central in evolution.
This is why even those who might be willing to acknowledge an organizing and coordinating capability, or agency, in individual development will nevertheless find the idea of such a capability impossible in the evolutionary case. What could conceivably coordinate all those communal interactions, including mating choices and predatorial patterns (and not forgetting mutational processes), so as to create a coherent evolutionary narrative? How could an individual organism “know” where its species happens to be headed in an evolutionary sense? And without knowing, how could it contribute meaningfully to the unfolding story?
We can agree: surely the relevant populations do not “know” where they are headed. But here again we can usefully recall the puzzle of development. For it is also true that the billions of cells in the embryonic horse do not “know” how to grow, differentiate, interact, and move coordinately toward the mature form and functioning of the adult animal. And yet — indisputably — the horse, in its whole being, does know where it needs to get to, and all its cells participate in that understanding.
For that matter, birds building their nests do not “know” about the needs of their unborn offspring, the individual wolf doesn’t “know” the collective strategy for bringing down an elk, and the hundreds of relevant molecules in a cell nucleus do not “know” how to coordinate their activity in order to repair a DNA lesion. Yet the relevant wisdom is in all cases visibly present and active, revealing itself as powerfully effective in development and behavior.
There is, however, a genuine question about how we are to understand the relation between intentional agencies observed at different biological levels. The relatively independent life of the cell is not the same as the life of the organism that integrates the activity of all its different cells. Similarly, the development of individual horses does not tell us all we would like to know about the intentions manifesting in the evolution from Eohippus to American Pharoah. It would be wrong to assume that the more or less fixed stages through which individual development passes give us a neat roadmap for the course of evolution.
Interpenetrating intentions. To speak, as in the preceding paragraph, of “different biological levels” is to be reminded that we confront different sorts of organic wholeness depending on the contextual breadth of our observation. We find ourselves looking at wholes embedded within still larger wholes. The cell is a functional whole, and so is the organ of which it is a part, and so again is the entire organism of which the organ is a part. But none of these is a whole in any absolute sense. If the “whole organism” were an absolute whole, entire unto itself, it would have no need to engage the world. Not belonging to the world, it would be unknowable by us.
The twentieth-century botanist, Agnes Arber, captured the relative character of organic wholes with her usual thoughtfulness:
The biological explanation of a phenomenon is the discovery of its own intrinsic place in a nexus of relations, extending indefinitely in all directions. To explain it is to see it simultaneously in its full individuality (as a whole in itself), and in its subordinate position (as one element in a larger whole).11
Every ecological setting, every organism within that setting, every organ within the organism, and every cell within the organ is a whole providing a context for its own interrelated parts, and at the same time is itself contextually embedded within larger wholes. “Context”, “whole”, and “part” can never be rigid, absolute terms in biology. They are bound up with interweaving spheres of activity. Certainly this is consistent with our experience of our own selves as more or less independent yet socially embedded beings.
This suggests that, however we may eventually come to understand the sources of biological, end-directed activity, we have no reason to associate centers of agency solely with “individuals” at any one level of generality. Just as our observation gives us interpenetrating wholes — from cells to organisms to populations and beyond — so, too, it gives us interpenetrating centers of agency. After all, what makes a whole a distinctive whole is its distinctive agency — the way in which, as a context, it “disciplines” its parts toward the realization of its own integral unity.
From flocks, herds, and schools, to bee and ant colonies, to parasitic and symbiotic pairs, to more or less closely aggregated communities of cells, to the highly differentiated and elaborately integrated cells of our own bodies, to earth’s biosphere as a whole — the one thing we can know directly is that we discover agency and intention wherever we see activity unfolding according to its own meanings. This includes purposeful, end-directed activity, whether we see it in bee colony behavior, or in the cells of a body, or in the facts of individual development, or in the beautiful evolutionary development of American Pharoah from Eohippus.
Evolution does, however, present practical problems to biologists that individual development does not. Cycles of development are endlessly and reliably repeated, so that no one can avoid at least implicitly acknowledging their end-directed character. Time and again, horse zygotes lead to adult horses. Evolution, by contrast, encompasses the totality of life on earth, and occurs only once. No more than in reading a good novel can we predict, mid-way through the story, its later outcome — just as, if we were viewing a tadpole for the first time, we could not predict its further development into a mature frog.12 The non-repeatability of evolution makes it all too easy, for those bent on doing so, to “forget” everything they know about the creative and end-directed character of all the life processes through which evolution occurs.
Evolution as a picture of creativity and coordination. To speak about the directed and future-oriented character of evolution is not to suggest that there is any simple, uniform, linear progress, traceable along a single line leading from ancestors to descendants. The picture of evolution that has long been coming into focus is one of great complexity and nonuniformity of change. Partly in response to environmental novelties, the creative roots of life “blossom” with an almost incomprehensible luxuriance and diversity of forms, producing forward glimpses here, throwbacks there, and extinctions almost everywhere. New developments may be foreshadowed in very different ways in different intersecting and diverging lineages, with the various, partially expressed potentials sometimes coming together and reaching their full fruition in a previously inconspicuous line.
Whole-organism biologist Craig Holdrege points out that even where we have a rich supply of fossils, as in the horse family, “there is a surprising diversity in the forms that predate modern horses”. And, regarding human fossil history, “the more fossils that are found, the less straightforward the emerging picture of the evolving human form becomes”:
If we consider this feature of the fossil record from a bird’s eye perspective, it is as if we are seeing hints of what is to come spread out in various earlier forms, which then become extinct. Eventually new forms appear, sharing characteristics with various earlier forms but in a new configuration that could never have been predicted on the basis of what came before.13
This nonlinear character of evolutionary “lineages” throws light on the problem of common ancestors (or “missing links”).14 The seemingly erratic pattern Holdrege summarizes makes it hard to see why we should expect to find unambiguous common ancestors of the sort we might like to imagine. It is a live question whether there are many — or any — uncomplicated, smooth pathways from progenitor types to substantially different descendent types, marked by a straightforward accumulation of new features, whereby we can witness a single, smooth, continuous line of transformation. Whatever the nature of the intention we see broadly at work, it must be a collective intention.
While some might take non-simple pathways of evolution as an argument against the role of intentional agency, the truth is just the opposite. For, as we have seen in the lives of individual organisms, it is precisely the purposive or intentional aspect that accounts for the ultimate coordination of otherwise more or less free and independent physical events. In this connection, we might think of evolution as embodying Weiss’ principle of micro-indeterminacy and macro-determinacy — only now writ very large indeed. What happens on a broader scale is not a simple determination or extrapolation predictable from the smaller scale, but is the result of a coordination of events toward living ends.
Or, again, there is Holdrege’s intentionally provocative and negative answer to the question, “Do Frogs Come from Tadpoles?” The tadpole is prerequisite for the frog — it provides necessary conditions for the frog — but the tadpole does not account for the frog. We do not find the frog as a necessary outcome “dictated” by the physical being of the tadpole. The metamorphosis shows us rather, the same creativity evident in all biological activity. What both the problem of missing ancestors and the widespread fact of organismal metamorphosis suggest is that we may as yet hardly have guessed how major evolutionary transformations are creatively coordinated.
I have heard it suggested that the human individual is more closely analogous to the typical species than to an individual of such a species. Whatever one thinks of that, the biographies of human individuals are one form of evolving life on earth, and they throw useful light on the kind of evolutionary process we have just now been considering. In a biography we are not at all surprised by all sorts of subplots, both productive and abortive, woven together in the most tortuous ways. Yet, when we examine the biography as a whole, we find that there is a story to be told — one that, as many excellent biographers have demonstrated so well, can be presented from diverse angles, while still holding together as one story. In other words, we are looking at neither a smooth, predetermined logical progression, nor a mere succession of events without coherence. Rather, we watch a playing out of interacting meanings, a kind of narrative tapestry.
What is missing in current evolutionary theory is not, to begin with, crucial pieces of new evidence (although, given the extent of our current ignorance about the most basic historical facts, we could certainly use a few of those), but rather a willingness to look at the narrative tapestry of life on earth with something like the biographer’s attentiveness.
We should be clear about the real sticking point for biologists. The fact that most of the cells in a tightly knit body are physically contiguous and therefore subject to certain physical causes does not in any relevant sense distinguish the working of biological intention in such a body from its working in evolutionary transformation. The organisms in an evolving population have no fewer causal connections than the cells in an individual. Eating and being eaten are surely causal! — a fact that, quite reasonably, figures centrally in conventional theory. And there is also the role of cognition. If, as many do today, we acknowledge a kind of cognition in cells sensing and responding to each other, how much more should we acknowledge the causal (not to mention the intentional) connections between all those organisms possessing specialized sense organs!
But while physical and chemical causal relations are certainly prerequisites for coherent transformation, whether in development or evolution, causal events do not explain their own coordination in extended living narratives. This is the crux of the matter. The reluctance of biologists to face the evident reality of evolution as a coherent narrative does not lie in the very real differences between development and evolution, but rather in a refusal to deal seriously with the problem of active biological wisdom and intention in either case.
This wisdom does not arise from physical and chemical interactions, but plays through them. We have no more reason to consider the guiding ideas at work in organisms as alien to their material being than we have for considering the mathematical ideas defining physical laws as alien to material being. If, in the one case, ideas come into play at the very root of physical manifestation (we do not have matter first, and then the lawful ideas it obligingly “obeys”), there is no reason to view the distinctive ideational content, or wisdom, governing organismal life in an altogether different way.
And so, just as we cannot separate matter from its ideal lawfulness, we also (as philosopher Ronald Brady has explained) “cannot detect, in [organic] phenomena, the distinction between ‘that which is to be vitalized’ and ‘that which vitalizes’”. And therefore we have no business “bother[ing] ourselves about how to add the potency of life to the stuff of life”15 — no more than we have any business worrying about how to add to matter the potency of the ideal and mathematically succinct law of gravity.
A few critical observations about current theory — especially the idea of fitness — may help to bring out the force of all the preceding considerations.
To begin with, no one has any clear or fully workable idea about what “fitness” means. Evolutionary biologists are required to assign numerical values to fitness — values enabling reasonable comparison of traits and organisms, so that we can determine which is fitter than which. But how do we take all the infinitely wide-ranging and interwoven considerations that might bear on fitness and reduce them to a single quantity? It is a practical impossibility. As a pair of philosophers put it in a 2005 article, “Suppose a certain species undertakes parental care, is resistant to malaria, and is somewhat weak but very quick. How do these fitness factors add up? We have no idea at all.”16
John Beatty, a major contributor to the most popular theory of fitness (a now rather shopworn and probabilistic theory broached in 1979 and known as the propensity theory), remarked in 1992 that “The precise meaning of ‘fitness’ has yet to be settled, in spite of the fact — or perhaps because of the fact — that the term is so central to evolutionary thought.”17
This is, if anything, even more emphatically true today. The concept remains troubled, as it has been from the very beginning, with little agreement on how to make it a workable part of evolutionary theory. Indeed, the “consensus view”, as Roberta Millstein and Robert Skipper, Jr., write in The Cambridge Companion to the Philosophy of Biology,18 is that “biologists and philosophers have yet to provide an adequate interpretation of fitness”.
And Harvard geneticist Richard Lewontin, along with his co-author, University of Missouri philosopher André Ariew, expressed the conviction that “no concept in evolutionary biology has been more confusing” than that of fitness.19 Yet, as two other philosophers of biology remind us, “the theory of natural selection as empirical science" hinges on what we mean when we say “x is fitter than y”.20
How do we account for the arrival of the fittest? Suppose it had turned out that there is no evolution — that, as was believed for millennia, each living “kind”, or species, remained faithful to a statically understood essential nature. It would still be true that some organisms (the “fittest”) would be most likely to survive and reproduce, with “defectives” presumably being selected for elimination. Such a world would in no way contradict the evolutionist’s idea of fitness, but would remain perfectly compatible with it.
This shows that neither the basic concept of fitness nor the closely allied concept of the “natural selection of the fittest” necessarily implies any sort of evolutionary transformation. Everything depends on what sort of beings organisms are, and what they are actually doing — in particular, what sort of fitness they might be moving toward. And if they are doing something transformative, then what they are doing is the decisive question. In other words, to focus on the general abstraction of fitness as such is simply to ignore the possibility that organisms might be in the business of “going somewhere” in an evolutionary sense.
The lack of explanatory value in the ideas of fitness and natural selection, when considered apart from the actual sources of transformation, has, in fact, long been recognized. The problem was given its most memorable statement by the Dutch botanist and geneticist, Hugo de Vries, who said during the first decade of the twentieth century, “Natural selection may explain the survival of the fittest, but it cannot explain the arrival of the fittest”.21
Or, as that preeminent zoologist of the middle twentieth century, Adolf Portmann, remarked in 1967: Regarding specific traits, natural selection “might afford a reason for their preservation, but never provide the cause for their origin”.22
And, in 2011, Lynn Margulis, a pioneer in exploring the role of symbiosis in evolution, said that “Natural selection eliminates and maybe maintains, but it doesn’t create”.23
Against this sort of objection, Stephen Jay Gould once asked, “Why was natural selection compared to a composer by Dobzhansky; to a poet by Simpson; to a sculptor by Mayr; and to, of all people, Mr. Shakespeare by Julian Huxley?” The answer, according to Gould, is that these allusions helpfully underscore the “creativity of natural selection”:
The essence of Darwinism lies in its claim that natural selection creates the fit. Variation is ubiquitous and random in direction. It supplies the raw material only. Natural selection directs the course of evolutionary change. It preserves favorable variants and builds fitness gradually.24
On its face, Gould’s argument was a puzzling one. He seemed to be saying that, because organisms are so expert and so prolific at producing new possibilities of life (“variation is ubiquitous”), we can ignore their creative achievement when we theorize about the historical genesis of new possibilities of life. Because organisms so abundantly provide raw materials for creative work, we are somehow free to declare a pure abstraction — natural selection — the agent performing the real biological work. It need only preserve the right collection of all those freely provided raw materials.
How easy it is, apparently, to forget that the “raw materials” are never merely raw materials! At the first appearance of any evolutionarily useful trait, or any modification of such a trait, the creative work has already been accomplished. We find ourselves looking, not at random raw materials, but at viable features harmoniously incorporated into living beings by the only power capable of such incorporation — a transformative power oriented toward the future while also resistant to whatever does not reflect the character and developmental potentials of its own dynamic type.
The concept of fitness conveniently substituted for the bother and difficulty of having to deal with the complexities (and philosophical distastefulness) of this creative work. The organism’s agency was now transferred to the supposedly creative “pressure” of natural selection, a “force” that acted on random (for example, “cosmic ray-induced”) mutations to either eliminate them or spread them through a population, based on their “fitness”.
The flow of mutated genes through a population had the decisive advantage that, in combination with a fuzzy concept of fitness, it offered some convenient rules of thumb that in turn yielded to mathematical treatment, unlike the organism’s actual creative organizing powers. These rules applied with rough (statistical) validity to the movement of genes within a population, and to this day discoveries and theorems about such movement are widely taken to be a direct demonstration of our knowledge of evolution. Genes just were, one assumed, the traits of evolutionary interest.
The credibility of this assumption — if there ever was any — is now long gone, even if many population geneticists have been extraordinarily slow in picking up on the relevant literature. And, in any case, the spreading of well-integrated new traits through a population does not explain the arrival of those traits in the first place. It explains nothing of the originating biological powers of evolutionary change.
The virtues of being unfit. As soon as we recognize the organism as an agent of transformation, we can see the limited relevance of the concept of fitness in another way. If organisms really are “going somewhere” — in this direction rather than that — then their supposed fitness may not only fail to explain the fact, but a fitness advantage is not even a general prerequisite for organisms contributing crucially to the success of the journey.
Individual development again provides a useful object lesson. Perhaps there are not many transformations greater than the one illustrated by frog metamorphosis. At the critical time of change, a tadpole is losing its organs for feeding on algae even as its organs for feeding on insects or other animals are being formed.25 And so also with many other organs and tissues. The animal’s overall condition during this period of intense change is awkward, perilous, and scarcely “fit”. But it is also the promise of something gloriously new.
The broader truth here is that the “weakest” organisms may at times be precisely those carrying the heaviest burden of the future. Transformation of any profound sort, by definition, entails the loss of past formation, along with consequent destabilization of the previous integral unity. In a coordinated process of fundamental change, we should expect that organisms barely able to survive might contribute decisively important heritable potentials to their offspring.
Actually, given phenomena such as lateral gene transfer and niche construction, along with the more general role of behavior in evolution, we can know that not even the ability to leave offspring is an absolute requirement for those organisms contributing to the evolution of their kind.
So perhaps instead of saying, “The most successful reproducers determine the future of the species”, we should say, “Those organisms representing the future of the species determine, over the long haul, what sort of individuals will become the most successful reproducers”. That illustrates the way evolutionary thinking will be turned inside out and backwards as soon as the obvious agency of organisms ceases to be ignored.
Everything I have written here — and critics may wish to take note — hinges on a single decision point: Are we willing to take seriously the meaningful and purposeful character of the life we witness in every organism? Can anyone claim that this is an issue wildly off the main path of biological science? Denis Walsh, one of our most insightful philosophers of biology, after noting that “organisms are fundamentally purposive entities”, expressed his perplexity by asking, “Why should the phenomenon that demarcates the domain of biology be off-limits to biology?”26 And the late, widely respected Chilean neuroscientist, Francisco Varela, wrote in his final paper: “The answer to the question of what status teleology should have in biology decides about the character of our whole theory of animate nature”.27
I have long thought: someone ought to look at what it might mean for evolution if we were to face up to these fundamental questions by taking organisms as we actually observe them, instead of pretending we do not see what is there. That is why I am writing the book so fragmentarily and cumbersomely summarized here.
If we are willing to inquire honestly whether evolution presents us with a meaningful narrative, then we must keep in mind that the later parts of any such narrative are often the key to understanding earlier parts. The unfolding reality comes to ever fuller expression. Whether watching a cat on the prowl, or tracing the unfolding life of a fertilized egg, or reading a human biography, we do not at first fully grasp what we are dealing with. As with all creative, future-oriented activity, it is only as future outcomes are clarified that we can recognize how they have informed the entire narrative.
When, then, we reflect upon the incredibly complex, end-directed tasks expertly carried out by vast collections of molecules even in the simplest and most primitive cells, it is natural to call to mind the eons of evolutionary transformation that have led from single cells to our own experience as conscious and willful agents pursuing our own meaningful tasks. Does the human outcome illuminate primordial origins?
It would, of course, be a fatal error to collapse all distinctions and talk about those early cells in the same way we talk about conscious human cognition and behavior. But the error would be equally egregious if we simply ignored the evident relation and historical continuity between the earliest forms of life and ourselves.
And, in fact, the evolutionary outcome does throw an intensely revelatory light upon the earlier circumstances. Our own bodies contain countless cells that function in many respects just like the most primitive cells we know. That is, they are engaged in meaningful, future-oriented activity — as, for example, in cell division — where part-processes are disciplined by their larger context. But, in our case, there is a still larger context whereby our highest capacities can, in some limited yet powerful respects, become the “guiding forces” of all that cellular activity.
Who will deny that our conscious intentions are carried out by our physiology, all the way down to the molecules in our cells? Everything shapes itself to our will, whether in the gentlest expressive movement of a pianist’s fingers, or the herculean effort of a shot putter. How could this not be a decisively important fact? — our organs, cells, and molecules capably embody and express our guiding intentions! And this truth is rendered perfectly natural by the fact that, quite apart from our conscious intentions, all our physiological activity shows, in its own right, a task-oriented, end-directed character.
In other words, our cellular activity, possessing a powerful ability to pursue intentional narratives, bends with exquisite sensitivity toward the realization of our intentions. Do we have any biological fact more fundamental than this to guide our scientific explorations?
We can hardly help asking: What are the guiding intentions in those many end-directed aspects of our physiology that are not now under our conscious control. And the question naturally carries over to the most primitive, single-celled forms of life we know, where we find the same well-coordinated and purposive functioning as in the cells of our own bodies. What are their guiding intentions?
The obvious next thing to ask at this point is whether our evolutionary origins can, with any reason at all, be thought of in the usual manner as mindless and meaningless. Or do those origins express high intentions that we humans, in consciously mastering our own bodies, and on our trajectory toward the future, have so far learned to touch only very lightly, being distracted as we now are by our abilities to toy mechanically with the exteriors of things?28
2. Quoted in Russell 1938, pp. 7-8. I have added paragraph breaks. The book by Nicholson is entitled How Birds Live: A Brief Account of Bird-Life in the Light of Modern Observation, and was published in London by Williams and Norgate, Ltd., in 1927.
4. While much of what I say will, with appropriate qualifications, be relevant to the plant kingdom and unicellular organisms, my references here to “organisms”, “living things”, and so on, should, for simplicity’s sake, be taken with animals specifically in mind.
5. The reference is to what is often referred to as “Laplace’s demon”, although Laplace himself did not use the word “demon”:
We ought then to regard the present state of the universe as the effect of its anterior state and the cause of the one which is to follow. Given for one instant an intelligence which could comprehend all the forces by which nature is animated and the respective situation of the beings who compose it — an intelligence sufficiently vast to submit these data to analysis — it would embrace in the same formula the movements of the greatest bodies of the universe and those of the lightest atom; for it, nothing would be uncertain and the future, as the past, would be present to its eyes … The curve described by a single molecule in air or vapour is regulated in a manner just as certain as the planetary orbits; the only difference between them is that which comes from our ignorance. (Laplace 1951, p. 4)
8. It is worth noting here the evidence for the fact that a human embryo also commonly takes in some cells from other organisms, particular the mother and the twin sibling, if there is one. But this does not disturb the general and universally agreed point I am making.
9. See, for example, Harvard geneticist Richard Lewontin (1992): “First, DNA is not self-reproducing, second, it makes nothing, and third, organisms are not determined by it. DNA is a dead molecule, among the most nonreactive, chemically inert molecules in the living world ... While it is often said that DNA produces proteins, in fact proteins (enzymes) produce DNA”.
10. There is every indication that this situation will inevitably change, as is already suggested by the current interest in (as well as resistance to) the idea of epigenetic inheritance.
28. A related question might be: “Do we live in a universe of beings, rather than things?” The answer to this question given during the past few hundred years, is, from a historical perspective, eccentric. Readers interested in such matters might want to read my article, “A Physicist, a Philologist, and the Meaning of Life: Do We Have a Home in the Vast Cosmos?”
Arber, Agnes (1985). The Mind and the Eye: A Study of the Biologist’s Standpoint, with an introduction by P. R. Bell. Originally published in 1954. Cambridge: Cambridge University Press.
Ariew, André and R. C. Lewontin (2004a). “The Confusions of Fitness”, British Journal for the Philosophy of Science vol. 55, no. 2, pp. 347-63. doi:10.1093/bjps/55.2.347
Beatty, John (1992). “Fitness: Theoretical Contexts”, in Keywords in Evolutionary Biology, edited by Evelyn Fox Keller and Elisabeth A. Lloyd. Cambridge MA: Harvard University Press, pp. 115-9.
Bouchard, Frédéric and Alex Rosenberg (2004). “Fitness, Probability and the Principles of Natural Selection”, British Society for the Philosophy of Science vol. 55, pp. 693-712. doi:10.1093/bjps/55.4.693
Brady, Ronald H. (1987). “Form and Cause in Goethe’s Morphology”, in Goethe and the Sciences: A Reappraisal, edited by F. Amrine, F. J. Zucker, and H. Wheeler, pp. 257-300. Dordrecht, Holland: D. Reidel. Available at https://natureinstitute.org/txt/rb/
DeVries, Hugo (1906). Species and Varieties: Their Origin by Mutation. Lectures delivered at the University of California, edited by Daniel Trembly MacDougal, second edition. Chicago: Open Court. Originally published in 1905. Full text available at https://gutenberg.org/ebooks/7234.
Gould, Stephen Jay (1976). “This View of Life: Darwin’s Untimely Burial”, Natural History vol. 85, pp. 24-30.
Holdrege, Craig (2017). Do Frogs Come from Tadpoles? Rethinking Origins in Development and Evolution. Nature Institute Perspectives #5. Ghent, NY and Great Barrington MA: Evolving Science Association. https://natureinstitute.org/pub/persp/5/index.htm
de Laplace, Pierre Simon, Marquis (1951/1814). A Philosophical Essay on Probabilities, translated from the sixth French edition by Frederick Wilson Truscott and Frederick Lincoln Emory with an introductory note by E. T. Bell.
Lewontin, R. C. (1992). “The Dream of the Human Genome”, New York Review (May 28), pp. 31-40.
Margulis, Lynn (2011). “Discover Interview: Lynn Margulis Says She’s Not Controversial, She’s Right”, Discover (April). Available online at "https://www.discovermagazine.com/the-sciences/discover-interview-lynn- margulis-says-shes-not-controversial-shes-right" "https://www.discovermagazine.com/the-sciences/discover-interview-lynn- margulis-says-shes-not-controversial-shes-right"
Matthen, Mohan and André Ariew (2005a). “How to Understand Causal Relations in Natural Selection: Reply to Rosenberg and Bouchard”, Biology and Philosophy vol. 20, pp. 355-64. https://link.springer.com/10.1007/s10539-005-5589-1
McDougall, William (1929). Modern Materialism and Emergent Evolution. New York: D. Van Nostrand.
Millstein, Roberta L. and Robert A. Skipper Jr. (2007). “Population Genetics”, in The Cambridge Companion to the Philosophy of Biology, edited by David L. Hull and Michael Ruse. Cambridge UK: Cambridge University Press.
Portmann, Adolf (1967). Animal Forms and Patterns: A Study of the Appearance of Animals, translated by Hella Czech. New York: Schocken.
Rich, Tina, Christine J. Watson and Andrew Wyllie (1999). Apoptosis: The Germs of Death, INature Cell Biology vol. 1 (July), pp. E69-71. doi:10.1038/11038
Russell, E. S. (1938). The Behaviour of Animals, second edition. London: Edward Arnold.
Talbott, Stephen L. (2014). “How Does the Organism Get Its Shape: The Causal Role of Biological Form”. Available at https://BiologyWorthyofLife.org/org/comm/ar/2014/brady_24.htm
Talbott, Stephen L. (2017). “Evolution and the Purposes of Life”, The New Atlantis # 51 (Winter). "https://thenewatlantis.com/publications/evolution-and-the-purposes-of- life" "https://thenewatlantis.com/publications/evolution-and-the-purposes- of-life"
Talbott, Stephen L. (2018). “A Physicist, a Philologist, and the Meaning of Life: Do We Have a Home in the Vast Cosmos?”. http://BiologyWorthyofLife.org/org/comm/ar/2018/meaning_33.htm
Walsh, Denis (2015). Organisms, Agency, and Evolution. Cambridge UK: Cambridge University Press.
Weber, Andreas and Francisco J. Varela (2002). “Life after Kant: Natural Purposes and the Autopoietic Foundations of Biological Individuality”, Phenomenology and the Cognitive Sciences vol. 1, pp. 97-125. Available at https://link.springer.com/article/10.1023/a:1020368120174
Weiss, Paul (1962). “From Cell to Molecule”, in The Molecular Control of Cellular Activity, edited by John M. Allen, pp. 1-72. The University of Michigan Institute of Science and Technology Series. New York: McGraw-Hill.
Weiss, Paul (1973). The Science of Life: The Living System — A System for Living. Mount Kisco NY: Futura Publishing.
Steve Talbott :: Evolution As It Was Meant To Be — An Overview