Good Enough: The Tolerance for Mediocrity in Nature and Society

good enough

The Tolerance for Mediocrity in Nature and Society

DANIEL S. MILO

Cambridge, Massachusetts

London, England

2019

Copyright © 2019 by the President and Fellows of Harvard College

All rights reserved

Cover image: Giraffe/Private Collection/Bridgeman Images

Cover design: Jill Breitbarth

978-0-674-50462-2 (alk. paper)

978-0-674-24005-6 (EPUB)

978-0-674-24006-3 (MOBI)

978-0-674-24004-9 (PDF)

The Library of Congress has cataloged the printed edition as follows:

Names: Milo, Daniel S. (Daniel Shabetaï), author.

Title: Good enough : the tolerance for mediocrity in nature and society /

Daniel S. Milo.

Description: Cambridge, Massachusetts : Harvard University Press, 2019. |

Includes bibliographical references and index.

Identifiers: LCCN 2018052620

Subjects: LCSH: Evolution (Biology) | Natural selection. | Social evolution. |

Imperfection.

Classification: LCC QH366.2 .M555 2019 | DDC 576.8—dc23

LC record available at https://lccn.loc.gov/2018052620

In memory of my beloved,

Naomi Aviv Milo,

who knew

Contents

Introduction

PARTONEIcons as Test Cases

1. The Giraffe: Science Begins in Wonder

2. The Domestication Analogy: Darwin’s Original Sin

3. The Galápagos and the Finch: Two Unrepresentative Icons

4. The Brain: Our Ancestors’ Worst Enemy

PARTTWOThe Theory of the Good Enough

5. Embracing Neutrality

6. Strange Ranges: The Bias toward Excess

7. Nature’s Safety Net

PARTTHREEOur Triumph and Its Side Effects

8. The Invention of Tomorrow

9. Humanity’s Safety Net

10. The Excellence Conspiracy: Critique of Evolutionary Ethics

Notes

Acknowledgments

Illustration Credits

Index

The world of things living … has room, wide but not unbounded, for variety of living form and structure, as these tend towards their seemingly endless but yet strictly limited possibilities of permutation and degree: it has room for the great and for the small, room for the weak and for the strong … The ways of life may be changed, and many a refuge found, before the sentence of unfitness is pronounced and the penalty of extermination paid.

—D’Arcy Wentworth Thompson, On Growth and Form (1942)

Introduction

Sigmund Freud wrote of three successive scientific outrages upon [humanity’s] naive self-love. The first was inaugurated by Copernicus, who discovered that our earth was not the center of the universe, but only a tiny speck in a world-system of a magnitude hardly conceivable. The last was Freud’s own: the revelation to the ego of each one of us that he is not even master in his own house. Between them was an idea that arrived upon the instigation of Charles Darwin, [Alfred Russel] Wallace, and their predecessors. This was the evolution of species by means of natural selection, which robbed man of his peculiar privilege of having been specially created, and relegated him to a descent from the animal world, implying an ineradicable animal nature in him.¹

The repercussions of two of those outrages were far-reaching. While the Copernican revolution resists popular appropriation—perhaps because we are not capable of experiencing the earth as other than flat and the sun as other than rising and setting—evolution and psychoanalysis mold our worldview. This despite the fact that few of us have opened The Interpretation of Dreams, and even professional biologists rarely read On the Origin of Species. Freud’s key concepts—subconscious; Oedipal complex; libido; reality and pleasure principles; defense mechanisms; sublimation; narcissism; id, ego, and superego—are embedded in daily life no less than in therapeutic practice.² Darwinism is similarly inescapable. Natural selection, struggle for survival, Malthusian competition, humanity’s apish origin, and adaptation are all pervasive in thinking about nature and human communities.

In particular, Darwinism courses through the ethics of capitalism. The latter’s terms—maximization, optimization, competitiveness, innovation, efficiency, cost-benefit trade-offs, rationalization—draw on the authority of Darwinian views of nature. Social Darwinism may be passé, but natural capitalism is fully alive. In the pages that follow, I will argue that Darwinism and neo-Darwinism—the mating of natural selection and genetics established in the mid-twentieth century—have a lot in common with neoliberalism. Nature knows what she is doing; in the market we trust. Homo economicus and animal economicus pursue the same goals and obey the same rules.

If evolutionary thought all too easily naturalizes capitalism’s competitive zeal, it is because Darwin was only partly right. There is no doubt that all organisms evolved from a common ancestor. But Darwin strayed in attributing evolution overwhelmingly to natural selection. It is this view that inspired Herbert Spencer, the father of social Darwinism, to coin survival of the fittest. At Wallace’s suggestion, Darwin himself eventually adopted this concept, too.³ It remains the basis of popular understanding of evolution today: everyone knows that nature is harsh and only the strong survive. But they are only partly right: nature is harsh sometimes; the strong survive often, but the weak stand a chance, too.

Both experts and the public join Darwin in overemphasizing natural selection, thereby distorting our sense of its role in both nature and humanity. Natural selection does occur, but there are nonadaptive mechanisms of change as well, such as genetic drift, geographic isolation, and the founder effect. None of these paths to survival rewards the hardest struggler or the best specimen. Their rewards are governed not by merit but by chance.

Genetic drift, the most prominent of these mechanisms, is the random change in the relative frequency of a gene variant, or allele, within a population. Genetic drift results in the survival not of the fittest but of the luckiest. One example is the northern elephant seal (Mirounga angustirostris). This seal experienced what evolutionary biologists call a bottleneck: an abrupt and drastic reduction in population size due to an environmental shock. By the late nineteenth century, the northern elephant seal had been harvested nearly to extinction for the oil derived from its blubber. Just a hundred or so specimens from Guadalupe Island, off Baja California, survived under the protection of the Mexican government. Those seals’ alleles persist in today’s population not because they were selected but because they got lucky.

Geographic isolation occurs when a population is cut off from the other members of its species. It can lead to speciation by means of inbreeding within a narrowed gene pool. The isolated population has only so many variants in it, and only these can be passed down and recombined in the next generation. The result is divergence from the original population, as each is now reproducing a different set of alleles. The origin of the new species is not natural selection but chance.

Likewise, the founder effect occurs when a few members of a population strike out from their habitat and start a new community. But though these founders may initiate an enduring lineage, they are not necessarily the fittest of their kind, nor do they necessarily adapt to their new environments through the fixation of advantageous mutations. A famous example is the small group of pigeons that landed on the island of Mauritius ages ago, lost their ability to fly, gained twenty kilograms, and evolved into Raphus cucullatus, also known as the dodo.⁴ The origin of this species was due to chance.

Specialists understand all this, so they avoid using survival of the fittest. And they know that natural selection does not cull every useless, exaggerated, and inefficient mutation, preserving only the best. Yet in 2007, Richard Dawkins defended the image of nature as "a miserly accountant, grudging the pennies, watching the clock, punishing the smallest extravagance. Unrelentingly and unceasingly."⁵ Indeed, evolutionary scientists as a group have done little to disabuse the public of Darwinian conceptions. Though the list of phenomena known to pass under natural selection’s radar is long and getting longer, scientists behave as though natural selection were natural law. They may not do so as openly as Dawkins, but their bias shows in their nearly exclusive focus on the discovery and confirmation of adaptations. Journalists, reflecting and reinforcing the public’s selection bias, trawl issues of Science and Nature for the latest evidence of possible adaptations, not the more complicated story of how evolution really works.

In that story, the humble and the humbled survive and multiply. The inefficient and the wasteful are certainly not the fittest, but they are good enough to make do. Natural selection has fashioned beautiful works, but it also tolerates a great deal of mediocrity. Even Darwin and Wallace thought so, before the Origin’s publication in 1859. Three years earlier, Darwin described nature as clumsy, wasteful, horridly blundering, the opposite of the grudging efficiency monger.⁶ That same year, Wallace affirmed nature’s tolerance for inutility: Do you mean to assert, then, some of my readers will indignantly ask, that this animal, or any animal, is provided with organs which are of no use to it? Yes, we reply, we do mean to assert that many animals are provided with organs and appendages which serve no material or physical purpose.⁷ This book aims to restore attention to what Darwin and Wallace perceived before their co-discovery committed them to hardened selectionism.

In addition, I hope to correct what are commonly understood as the prescriptive ramifications of evolutionary theory. Despite official disgust with Spencer and eugenics, survival of the fittest continues to underwrite political principles. We see the notion of Darwinian competition at the foundation of a merciless meritocracy. The winner owes survival / success to excellence, and the loser owes extinction / failure to its absence. The former has only him- or herself to congratulate; the latter, only him- or herself to blame.

As I detail in Chapter 2, Darwin himself first made this leap from nature to society. "All organic beings are striving to seize on each place in the economy of nature," he wrote in the Origin. If any one species does not become modified and improved in a corresponding degree with its competitors it will be exterminated.⁸ In the years since, biology has stamped its seal of approval on market fundamentalism. Survival of the fittest is gospel in the thinking of Milton Friedman, per Robert Frank’s The Darwin Economy: Liberty, Competition, and the Common Good.⁹ The business press tells us that economic cycles are Darwinian, picking off weak companies and leaving survivors stronger.¹⁰ Investment firms preach the doctrine of corporate Darwinism.¹¹

Whether we should, in the first place, look to nature for social models is an important and complex question but one I don’t address. Instead, I only wish to propose that nature cannot be called upon to endorse capitalist ethics because it prefers inertia and copy-paste to change and originality. The replication machinery of the DNA aims at a 100 percent success rate, with each new generation a perfect likeness of the previous. Organisms migrate only under stress; they would much rather stay home. There is, in general, no better tactic than to imitate one’s progenitors, since they have already proved their talents in the arts of survival and reproduction. Just ask the coelacanth (Latimeria chalumnae). This lungfish, known as the living fossil, has barely changed during the last four hundred million years.

In nature, change is either an accident, as in mutations, or a last resort, as in migration. It is never an end in itself. In the human society modeled on supposed Darwinian truths, change may still be motivated by accident or necessity, but change for change’s sake is all around us. Stagnation is the antithesis of all that capitalism and its culture demand, the heresy banished from the temple of the God Growth. Rather than rest, we are programmed to need new products, new artworks, new research. Under the regime of planned obsolescence, innovation follows its own urges, not the failure of old models. Industry, academia, politics, fashion—all modern economic and cultural efforts create products with artificially limited life cycles.

As a result, most of what we produce and consume is excess. Our so-called needs usually have nothing to do with our survival. Humans can do with less, a lot less, of almost everything, from surgical specialties and breeds of dog to varieties of breakfast cereal and synonyms for wonderful. And while we may at times struggle for existence, some of us more than others, most days are mostly peaceful, with little need of competition to prove our worth. We know this because, with all due respect to Friedman, there is such a thing as a free lunch. As I discuss in Chapter 9, human communities do a great deal to care for their members at no charge. This book is an example of what can be obtained gratis; it would have been impossible without the help of numerous biologists who charged neither money nor credit for their counsel.

Human society is not ruthlessly competitive, and neither is nature. Both are tolerant of excess, inertia, error, mediocrity, and failed experiment. Where great successes occur in society and in nature, luck can be far more important than talent. Yet there are many who tell us that talent—sometimes rendered as fitness, sometimes as merit—is all that matters in nature and in human affairs, each following a deep Darwinian law of the universe. This is the dogma I seek to undermine.

Key Scientific Concepts

Before questioning Darwinism in greater detail, I have to specify what I mean by that term, because in fact two theories are found under its umbrella: descent with modification, sometimes referred to simply as evolution, and natural selection. According to the former, Darwin wrote in the Origin, Probably all the organic beings which have ever lived on this earth have descended from some one primordial form, into which life was first breathed.¹² Darwin illustrated this prophetic speculation with a tree of life showing new species branching from predecessors, some continuing and others dying off (Figure I.1). Molecular biology proves Darwin right. The theory of natural selection holds that descent with modification is directed by a process "daily and hourly scrutinising, throughout the world, every variation, even the slightest; rejecting that which is bad, preserving and adding up all that is good; silently and insensibly working, whenever and wherever opportunity offers, at the improvement of each organic being in relation to its organic and inorganic conditions of life."¹³ This book argues that Darwin was sometimes right and often wrong.

Darwin viewed descent with modification and natural selection as inextricably linked, hence the full title of his On the Origin of Species by Means of Natural Selection, or The Preservation of Favoured Races in the Struggle for Life. This pairing of two distinct theories became orthodoxy, though it faced skepticism in Darwin’s own time. The common ancestry of all extant organisms was received as such by many of Darwin’s contemporaries; it was natural selection that initially received pushback from fellow evolutionists. St. George Jackson Mivart, a nineteenth-century disciple, bought into descent with modification but rejected natural selection entirely.¹⁴ Darwin took Mivart seriously enough to dedicate a chapter of the Origin’s sixth and last edition, Miscellaneous Objections to the Theory of Natural Selection, to his rebuttal. Another acolyte, George Romanes, did not reject natural selection but questioned its sole authorship of evolution. I have found it impossible to entertain a doubt, either upon Evolution as a fact, or upon Natural Selection as a method, Romanes wrote. That is, natural selection is not a fact but one way to look at facts. Romanes pointed to other possible sources of evolution, such as speciation through geographical isolation, now called allopatric speciation.¹⁵ Throughout this book, I return to Romanes to emphasize his underappreciated yet visionary contributions.

Figure I.1. Darwin’s Tree of Life: The Origin of Species illustrates descent with modification, showing numerous lineages emerging, persisting, and dying out across fifteen generations. Every species is a product of descent with modification, but Darwin erred in asserting that natural selection is overwhelmingly the agent underlying this process.

Another contemporaneous objection came from Wallace. The co-discoverer of natural selection was its most ardent defender, yet even he granted special dispensation to Homo sapiens in a paper titled The Limits of Natural Selection as Applied to Man.¹⁶ Wallace attributed human evolution to a WILL of higher intelligences or of one Supreme Intelligence. That Wallace erred in the direction of intelligent design avant la lettre speaks to the blurred boundary between that false idea and Darwin’s overzealous selectionism, as I discuss later in this introduction and in Chapter 10.

Much as there are two theories under the heading of evolution, we might also say that there are two evolutions. The first, celebrated by the title On the Origin of Species by Means of Natural Selection, accounts for speciation or interspecific variability. This is evolution as Darwin and his successors understood it. The second evolution is concerned with the existence of a species between its origin and its extinction; its normalcy, if you wish. The average lifespan of a species is between one million and ten million years. From the standpoint of Darwinian evolution, this time is uninteresting because it is static. But, in fact, during this time, there is much change; species accumulate variability, both qualitative and quantitative, though it may have no adaptive impact. The theory of the good enough aims to account for this intraspecific diversification.

What I am suggesting is that during stasis, organisms have the freedom to vary unimpeded and unimproved by natural selection. During certain explosive moments in natural history, environmental events and forced migration have obliged organisms to adapt to new niches—new competitors and changed habitats—or die. This is the first evolution. Those that do adapt pass along traits that make their descendants more robust and therefore more impervious to factors producing extinction. These descendants spend most of their lives at relative peace, so they can accumulate neutral variants—that is, variants that confer no benefit. There are enough resources to go around, and the fundamentals of life are sufficiently strong that any variant—excellent or mediocre—can survive as long as it isn’t lethal.

Biologists do appreciate the concept of neutral mutation, which was introduced in the 1960s by the population geneticist Motoo Kimura. But this theory is limited to the genotype; it does not concern phenotypic expression or behavior. Phenotypic neutrality, which I propose, is a Darwinian oxymoron. A phenotype that is not beneficial to its bearer is detrimental by definition, since it costs energy but earns nothing in exchange. Therefore, when faced with apparently useless or deleterious traits, biologists assume either that the traits are beneficial but for reasons not yet discovered or that natural selection will eventually catch up with them.

Accepting tolerance for neutrality at the phenotypic level would be a paradigm shift. Darwin seemed to leave open the possibility, but with considerable reservation. In The Descent of Man, he wrote, I did not formerly consider sufficiently the existence of structures, which, so far as we can at present judge, are neither beneficial nor injurious; and this I believe to be one of the greatest oversights as yet detected in my work.¹⁷ Much faith is invested in that at present—we may not yet know the benefits of a given trait, but someday we will. Regardless, Darwin’s followers never gave these musings much credence. Romanes, however, was an exception. A very large proportion, if not the majority, of features which serve to distinguish species from species are features presenting no utilitarian significance, he wrote.¹⁸ Yet his position all but disappeared from biology after the 1890s, overwhelmed by a view more in line with Wallace’s hardened selectionism: "None of the definite facts of organic nature, no special organ, no characteristic form or marking, no peculiarities of instinct or of habit, no relations between species or between groups of species, can exist but which must now be or once have been useful to the individuals or races which possess them."¹⁹

Against the heavy weight of tradition, I follow Romanes and extend the principle of neutrality to all strata of life, from the molecular to the behavioral. Granted, proving uselessness is as impossible as proving that you don’t have a sister. Instead, I approach uselessness via a close cousin: excess. What is excess? When you can do the same, or better, with less. What is uselessness? When you can do without.

The orthodox position holds that quantitative excess will be culled because it imposes metabolic burden: having more of something than needed forces an organism to do the hard work of obtaining additional calories, which is an adaptive disadvantage under conditions of Malthusian competition. So, to the extent that excessive traits are present in nature, we should doubt the role of natural selection in fixing and propagating them. As it turns out, having too much is almost as common in nature as it is in human society. For reasons I detail in Chapters 6 and 7, nature is strongly biased toward excess.

The distinction between qualitative and quantitative excess is critical. Even if we grant that all traits are or were once selected, the same is not necessarily true for their size and amount. For instance, giraffes cannot do without legs, but they would be better off with shorter ones in at least one respect: they give birth standing up, to newborns weighing seventy kilograms, who then fall head first.²⁰ Or consider the avocado. The average avocado tree has a million flowers but yields just one hundred fruits.²¹ Somehow avocado trees have managed to survive despite their profligacy. And one would be hard-pressed to come up with a selective explanation for the genomic excess of Paris japonica. This little flower has fifty times as many DNA base pairs as Homo sapiens.

If the prevalence of quantitative excess in nature is not enough to convince us of natural selection’s limits, then wide phenotypic ranges should be. It is an obvious but little acknowledged fact that most traits are viable across a range of amounts and sizes. For instance, a study of human kidneys in five racial groups from three continents found that people may have as few as 210,332 nephrons per kidney and as many as 2,702,079, with the healthy minimum located near the bottom of the range.²² There is no way natural selection chose nearly 3 million nephrons when about 500,000 would suffice. Wide ranges and optimization are incommensurable. Such ranges, the subject of Chapter 6, are smoking-gun evidence for natural selection’s chronic fallibility.

Wide ranges speak to what I call ontological neutrality. It simply does not matter whether one has 500,000 nephrons per kidney or 2 million. But as I discuss in Chapter 5, neutrality is not just a reality; it is also a way of looking at reality. I call this methodological neutrality. In criminal law, for instance, methodological neutrality is inherent in the presumption of innocence: everyone is innocent by default, and the burden of proof weighs on the accuser. In science, methodological neutrality is expressed by the null hypothesis, namely that every relationship between phenomena is, by default, the fruit of chance. The burden of refutation weighs on the researcher. You don’t have to prove innocence or justify chance.

Evolutionary biologists invert the principle: being selected is the default state, and a chance result is the outlier. There is a presumption of selection in nature, so a biologist is exempt from proving it. Instead, the burden of proof lies on whoever claims that a trait or a size was not selected. The time is ripe for biologists to embrace the ways of their fellow scientists and accept the null hypothesis: chance, which is to say, neutrality. Doing so does not imply rejection of natural selection, an indefensible stance. What it does imply is the rejection of natural selection as the default state in nature.

But while we cannot reject natural selection, we should recognize that, in the Darwinian mode, it is conceptually problematic. The trouble is that natural selection arises from a false analogy to artificial selection. As I discuss in Chapter 2, Darwin theorized natural selection in the image of a breeder who chooses one or another desirable trait from his specimens and promotes them in subsequent generations. We have this analogy to thank for the mistaken view that nature selects and that it selects only the best. This flaw in Darwinism was also known in the time of its creator. Wallace, though he became a devout selectionist, contested the domestication analogy as a major weakness in Darwin’s reasoning.

The domestication fallacy is to blame for the common misconception that nature is a subject that takes action. Nature doesn’t select, operate, cause, scrutinize, or purify. It doesn’t do anything. Rather, nature is a shifting set of conditions under which different traits make greater and lesser contributions to an organism’s probability of survival and reproduction. Very poorly adapted organisms will almost certainly be culled; their survival probability is virtually zero. Better-adapted organisms have a greater probability of surviving and reproducing. Their traits need not make them excellent, as wide ranges and excess demonstrate. Whether or not a species or an organism is well adapted, nothing has selected its traits. All we know is that these species and organisms are good enough not to die off. For this reason, I suggest substituting for natural selection the language of natural elimination or, better still, natural probability. However, to maintain continuity with the field of evolutionary biology, I typically use the consensual term natural selection to denote the process scientists have in mind when they speak of it.

Although biologists everywhere bemoan the language of nature as subject, practically all of them, and the public in their wake, remain loyal to the view that a breeder-like, optimizing natural selection is the default state of nature. How does natural selection maintain its dominant place in evolutionary thought despite its low relative frequency? I argue that the culprit is the biased human brain. It has a pronounced preference for the exceptional, which natural selection surely is, in the senses of being both rare and awe inspiring. One cannot but be impressed by the workings of, for example, adaptive selection. The Galápagos finches are the famed case, discussed in Chapter 3. All evolved from one species that drifted away from South America three million years ago. The finches descended from a generalist species, but having landed on islands with sparse resources, they had to develop specialties to survive. Thus, the cactus finch (Geospiza scandens) possesses a long beak with a