Category Archives: Philosophy

Science (Indeed, the World) Needs Fewer, Not More, Icons.

by Kenneth W. Krause.

Kenneth W. Krause is a contributing editor and “Science Watch” columnist for the Skeptical Inquirer.  Formerly a contributing editor and books columnist for the Humanist, Kenneth contributes frequently to Skeptic as well. He can be contracted at

The Sims statue and its protestors.

To the extent we are rational, we share the same identity.—Rebecca Goldstein.

September was an awkward month for Nature, perhaps the most influential and well-respected science publication on the planet.  In August, a group peacefully protested, and vandals subsequently defaced, a Central Park statue of J. Marion Sims, a 19th-century surgeon and founder of the New York Women’s Hospital often referred to as the “father of modern gynecology.”  Sims’ likeness was left with fiendish red eyes and the word “RACIST” scrawled across its back.

The quarrel stemmed from the mostly undisputed facts that, although Sims helped develop life-saving surgical techniques to help women recover from particularly traumatic births, he also experimented on female slaves without providing anesthesia, and after seeking consent only from their owners. Unsurprisingly, commentators contest whether Sims’ methods were consistent with the customs and scruples of his time (Washington 2017).

Nature’s first inclination was to publish an editorial originally titled, “Removing the Statues of Historical Figures Risks Whitewashing History,” arguing that we should leave such icons in place to remind passers-by of the important historical lessons they might provide (The Editors 2017). The piece also recommended the installation of additional iconography to “describe the unethical behavior and pay respect to the victims of the experimentation.”

Given then-recent events in the ever-emotionally explosive and divisive world of American popular culture especially, vigorous dissent was inevitable. A flurry of indignant letters descended on Nature’s editors.  Several writers suggested that, at least in America, the primary if not sole purpose of public statuary is to honor its subjects, not to inform curious minds of their historical significances (Comment 2017).  One contributor noted that the history of Nazi Germany has been well-documented in the very conspicuous absence of Nazi iconography.  Another reasoned that because written documentation always precedes statuary, removal of monuments would have “no impact on our understanding of the historical failings of those individuals.”

Other letters offered less restrained and, frankly, less disciplined commentary. One author submitted that the editorial “perpetuate[d] racist white supremacy.”  Two more branded it simply as “white privilege at its height” and as a “racist screed.”  Another found the article in support of “unethical science” and to inform Nature’s minority readers that they “remain unwelcome in science because of their race.”

Vandals defaced the Sims likeness with red paint.

But more importantly for my purposes here, many writers contributed thoughts on the Sims monument itself that reveal quite plainly our human tendencies to interpret the inherent ambiguity of statues—indeed iconography and other symbolic expressions more generally—consistent with our fears, personal agendas, or ideological mindsets. One author, for example, confided that the Sims statue bid her to “Go away, woman.  You have no authority here,” and to “Go away, woman of African descent.  You cannot have the intellect to contribute to the science of your own healthcare” (Green 2017).  Another saw Sims’ likeness as a “signal” that the “accomplishments of a white man are more important than his methods or the countless people he victimized,” and that “the unwilling subjects of that research … are unimportant and should be washed away.” (Gould 2017; Comment 2017).  Yes, all of that from a motionless, voiceless sculpture.

In the end, Nature’s guests called consistently for the icon’s swift removal.  And given its and any other statue’s essential ambiguity, I agree.  Take it away, melt it down, and donate its metal to a more fruitful purpose.  But, regrettably, many writers also petitioned for additional iconography—this time to honor accomplished females in medicine and the victims of sexist and racist medical practices.  In other words, they would display more monuments of more humans, no doubt all with potentially hideous skeletons lurking in their so far sealed closets, likely to be scrutinized and challenged by any conceivable number of equally fault- and agenda-ridden human interpreters to come.

In the rush to colonize others’ minds, or perhaps to cast painful blows against cross-cultural enemies, has anyone actually taken the time and effort to think this through? Both duly and thoroughly reproved, Nature’s editors quickly apologized and revised their article, including its title, to comply with reader objections (Campbell 2017; The Editors 2017).  But glaring similarities between the Sims controversy and more widely publicized events involving statues of Confederate generals, for example (at least one of which resulted in meaningless violence), have attracted the attention of the general media as well.

Police protect Charlottesville’s statue of General Lee.

Writing for The Atlantic, Ross Anderson aptly observed that “the writing of history and building of monuments are distinct acts, motivated by distinct values” (Anderson 2017).  No serious person ever suggested, he continued, that statuary “purport[s] to be an accurate depiction of its history.”  So far, so good.  At that critical point, Anderson appeared well on his way to advancing the sensible argument that inherently simplistic and ambiguous iconography can only divide our society, and perhaps even inspire (more) pointless violence.

Unfortunately, that was also the point where the author stumbled and then strayed onto perhaps well-worn, but nevertheless unsustainable trail. The legitimate purpose of a society’s statuary, he argued, is “an elevation of particular individuals as representative of its highest ideals,” a collective judgment as to “who should loom over us on pedestals, enshrined in metal or stone ….”  But, honestly, no credible history has ever instructed that any individual, no matter how accomplished, whether male or female, black or white, can ever represent our “highest ideals.”  And is there anything about recent American history to suggest we could ever agree on what constitutes those ideals?  And, come to think of it, how do people tend to react when others choose which monuments and symbols will “loom over” them?  Indeed, wasn’t that the problem in Charlottesville, Virginia?

White supremacists march on Charlottesville.

According to Anderson, the activists demanding removal of the Sims statue and its replacement with iconography of presumptively more deserving subjects ask only “that we absorb the hard work of contemporary historians … and use that understanding to inform our choices about who we honor.” But, as any experienced historian knows, historical facts can be, and often are, responsibly parsed and interpreted in many different ways.  And why should common citizens blindly accept one credible historian’s perspective over that of any other?  Regardless, shouldn’t we encourage the public to consult the actual history, rather than convenient, but severely underdeveloped and necessarily misleading shortcuts?

Author Dave Benner argued, instead, that we should preserve our monuments (Benner 2017). Pointing to the New Orleans statue of Franklin Roosevelt, which, to this point, remains free of public derision and vandalism, Benner reminded us of Executive Order 9066, by which FDR displaced 110,000 American citizens of Japanese ancestry into internment camps, without due process, in “one of the saddest and most tyrannical forms of executive overreach in American History.”  Should the FDR monument (indeed, the dime) be purged according to the same reasoning offered by Nature’s revised editorial and those who oppose the Sims statue?  By such a standard, would iconography depicting any of the American founders survive?

Perhaps not. But to what supposedly disastrous end?  By Benner’s lights, the removal of cultural iconography would “simply make it harder for individuals to learn from the past.”  But, again, as the many dissenter’s to Nature’s original editorial observed, the purpose of statuary is not to inform.  And let’s be completely candid here: nor is it to “honor” the dead and insensible subjects of such iconography who no longer hold a stake in that or any other outcome.  Rather, the unspoken object is no less than to decree and dispense value judgments for the masses.

And some would no doubt argue the propriety of that object in the context of politics and government. But can and should science do better?  “As the statues and portraits of Sims make clear,” offers Harriet Washington, award-winning author of Medical Apartheid, “art can create beautiful lies” (Washington 2017).  “To find the truth,” she advises, “we must be willing to dig deeper and be willing to confront ugly facts.  No scientist, no thinking individual, should be content to accept pretty propaganda.”

Science’s battle is not with any particular ideological foe. It stands against all ideologies equally.  It has no interest in turning minds to any individual’s, or any coalition’s social cause because it has no agenda beyond the entire objective truth.  Science is incapable of pursuing ambiguity or any shortcut, especially where the potential for clarity, completion, and credibility persists.  And science certainly doesn’t need more icons; it needs fewer, or none.


A final thought on symbolic expression:

Yes, American history is saturated with political symbolism, from the flags of the colonial rebellion to the Tinker armbands and beyond.  As I wrote this column, however, the discussion of alleged “race” in America grew increasingly inane—dominated, in fact, by Donald Trump, our Clown in Chief, on one side, and mostly mute and under-studied NFL football players on the other.  The social, popular, and activist media, along with their rapacious followers, of course, seemed thoroughly enchanted by this absurd spectacle.

I take no position on this “debate,” if it can be so characterized. Indeed, comprehension of the contestants’ grievances is precluded by their irresponsible methods.  The President’s very involvement is inexplicable.  But, for me, it’s the players’ exclusively symbolic expressions that cause greater concern.  Again, not because I disagree with whatever they might be trying to say.  Rather, because their gestures are so ambiguous and amenable to any number of conceivable interpretations that, in the end, they say nothing.  Is this the future of all public discourse?

Waving or burning flags just isn’t impressive. Nor is standing, or sitting when others stand.  Nor is raising a fist or locking arms.  Because these expressions require no real investments, they amount to cheap, lazy, conveniently vague, and, thus, mostly empty gestures.  I’m old enough to know that they’ll persist, of course, and no doubt dominate the general public’s collective consciousness.  I only hope we can manage to maintain, perhaps even expand, spaces for more sober, motivated, and responsible discourse.  In any case, I’d prefer not to spend my remaining years watching them being torn down, especially from within.



Anderson, R. 2017. Nature’s Disastrous ‘Whitewashing’ Editorial. Available online at; accessed September 27, 2017.

Benner, D. 2017. Why the Purge of Historic Monuments Is a Bad Idea. Available online at; accessed September 27, 2017.

Campbell, P. 2017. Statues: an editorial response. Nature 549: 334.

Comment. 2017. Readers Respond to Nature’s Editorial on Historical Monuments. Available online at; accessed September 26, 2017.

Gould, K.E. 2017. Statues: for those deserving respect. Nature 549: 160.

Green, M.H. 2017. Statues: a mother of gynaecology. Nature 549: 160.

The Editors. 2017. Science must acknowledge its past mistakes and crimes. Nature 549: 5-6.

Washington, H. 2017. Statues that perpetuate lies should not stand. Nature 549: 309.


CRISPR-Cas9: Not Just Another Scientific Revolution (Special Report).

by Kenneth W. Krause.

Kenneth W. Krause is a contributing editor and “Science Watch” columnist for the Skeptical Inquirer.  Formerly a contributing editor and books columnist for the Humanist, he writes regularly for Skeptic magazine as well.  He may be contacted at

Poised to transform the world as we know it, a new gene-editing system has bioethicists wringing their hands, physicians champing at the bit, and researchers dueling with demons.


Is it possible to overstate the potential of a new technology that efficiently and cheaply permits deliberate, specific, and multiple genomic modifications to almost anything biological? What if that technology was also capable of altering untold future generations of nearly any given species—including the one responsible for creating it?  And what if it could be used, for better or worse, to rapidly exterminate entire species?

Certain experts have no intention of veiling their enthusiasm, or their unease. Consider, for example, biologist David Baltimore, who recently chaired an international summit dedicated primarily to the technology’s much-disputed ethical implications.  “The unthinkable has become conceivable,” he warned his audience in early December.  Powerful new gene-editing techniques, he added, have placed us “on the cusp of a new era in human history.”

If so, it might seem somewhat anticlimactic to note that Science magazine has dubbed this technology its “Breakthrough of the Year” for 2015, or that its primary developers are widely considered shoo-ins for a Nobel Prize—in addition, that is, to the US$3 million Breakthrough Prize in Life Sciences already earned by two such researchers.  All of which might sound trifling compared to the billions up for grabs following imminent resolution of a now-vicious patent dispute.

Although no gene-editing tool has ever inspired so much drama, the new technology’s promise as a practical remedy for a host of dreadful diseases, including cancer, remains foremost in researchers’ minds. Eager to move beyond in vitro and animal model applications to the clinical setting, geneticists across the globe are quickly developing improved molecular components and methods to increase the technology’s accuracy.  In case you haven’t heard, a truly profound scientific insurrection is well underway.

Adapting CRISPR-Cas9.

Think about a film strip. You see a particular segment of the film that you want to replace.  And if you had a film splicer, you would go in and literally cut it out and piece it back together—maybe with a new clip.  Imagine being able to do that in the genetic code, the code of life.—biochemist Jennifer Doudna (CBS News 2015).

Genetic manipulation is nothing new, of course. Classic gene therapy, for example, typically employs a vector, often a virus, to somewhat haphazardly deliver a healthy allele somewhere in the patient’s genome, hopefully to perform its desired function wherever it settles.  Alternatively, RNA interference selects specific messenger RNA molecules for destruction, thus changing the way one’s DNA is transcribed.  Interference occurs, however, only so long as the damaging agent remains within the cell.

Contemporary editing techniques, on the other hand, allow biologists to actually alter DNA—the “code of life,” as Doudna suggests—and to do so with specific target sequences in mind.  The three major techniques have much in common.  Each involves an enzyme called a programmable nuclease, for example, which is guided to a particular nucleotide sequence to cleave it.

Then, in each case, the cell’s machinery quickly repairs the double-stranded break in one of two ways. Non-homologous end joining for gene “knock out” results when reconstruction, usually involving small, random nucleotide deletions or insertions, is performed only by the cell.  Here, the gene’s function is typically undermined.  By contrast, homology-directed repair for gene “knock in” occurs when the cell copies a researcher’s DNA repair template delivered along with the nuclease.  In this case, the cleaved gene can be corrected or a new gene or genes can be inserted (Corbyn 2015).

But in other ways, the three editing techniques are very distinct. Developed in the late 1990s and first used in human cells in 2005, zinc-finger nucleases (ZFN) attach cutting domains derived from the prokaryote Flavobacterium okeanokoites to proteins called zinc fingers that can be customized to recognize certain three-base-pair DNA codes.  Devised in 2010, transcription activator-like effector nucleases (TALENs) fuse the same cutting domains to different proteins called TAL effectors.  For both ZFN and TALENs, two cutting domains are necessary to cleave double-stranded DNA (Maxmen 2015).

The third and most revolutionary editing technique, and subject of this paper, consists of clustered regularly interspaced short palindromic repeats (CRISPR) and a CRISPR-associated protein-9 nuclease (Cas9). Introduced as an exceptionally precise editing technique in 2012 by Doudna at the University of California, Berkeley, and microbiologist Emmanuelle Charpentier at the Max Planck Institute for Infection Biology in Berlin, CRISPR-Cas9 is actually the bacterium Streptococcus pyogenes’ adaptive immune system that confers resistance to foreign elements, like phages and plasmids.


CRISPR thus refers to short bits of DNA seized from invading viruses and stored in the bacterium’s own genome for future reference, and Cas9 is the enzyme S. pyogenes uses to cleave a subsequent invader’s double helix.  In other words, in its native setting, CRISPR-Cas9 is the system a certain bacterium uses to recognize and disable common biological threats.  Unlike ZFN and TALENs, CRISPR-Cas9 does not rely on the F. okeanoites cutting domain and, as such, can cleave both strands of an interloper’s double helix simultaneously with a single Cas9 enzyme.

But what makes the CRISPR system so special, in part, and so adaptable to the important task of gene-editing, is its relative simplicity. Only three components are required to achieve site-specific DNA recognition and cleavage.  Both a CRISPR RNA (crRNA) and a trans-activating crRNA (tracrRNA) are needed to guide the Cas9 enzyme to its target sequence.  What Doudna and Charpentier revealed six years ago, however, were the seminal facts that an even simpler, two-component system could be developed by combining the crRNA and tracrRNA into a synthetic single guide RNA (sgRNA), and that researchers could readily modify a sgRNA’s code to redirect the Cas9 enzyme to almost any preferred sequence (Jinek et al. 2012).  Today, a biologist wanting to edit a specific sequence in an organism’s genome can quickly and cheaply design a sgRNA to match that sequence, order it from a competitive manufacturer for US$65 or less, and have it delivered in the mail (Petherick 2015).

None of which is to suggest that a CRISPR system is always the best tool for the gene-editing job, at least not yet. Critically, CRISPR-Cas9 is relatively easy to program and remains the only technique allowing researchers to “multiplex,” or edit several genomic sites simultaneously.  But TALENs have the longest DNA recognition domains and, thus, tend so far to result in the fewest “off-target effects,” which occur when nucleotide sequences identical or similar to the target are cut unintentionally.  And ZFNs are much smaller than either TALENs or CRISPR-Cas9, especially the most popular version derived from S. pyogenes, and are therefore more likely to fit into the tight confines of an adeno-associated virus (AAV)—currently the most promising vector for the delivery of gene-editing therapies.

Even so, CRISPR research continues to progress at breakneck speed. In 2014, the number of gene-editing kits ordered from Addgene, a supplier based in Cambridge, Massachusetts, for research using ZFN and TALENs totaled less than 1000 and less than 2000, respectively.  During that same year—only two years after the new technology was introduced, the number of kits ordered for CRISPR research totaled almost 20,000 (Corbyn 2015).  More importantly, rapidly increasing orders seem to have translated into significant results.  As 2015 ended and a new year began, new studies announcing the creation of smaller guide RNAs and, especially, the reduction of off-target effects began to dominate science headlines.

Building a Better Mousetrap.

At some point everyone needs to decide how specific is specific enough. The idea that you would make a tool that has absolutely no off-target effects is a little too utopian.—bioengineer Charles Gersbach (Ledford 2016).

It’s cheap, easy to use, and remarkably efficient, but CRISPR-Cas9 is not perfect. In early experiments, in fact, pathologist Keith Joung at the Massachusetts General Hospital in Boston, discovered that his enzymes were cutting unintended as often as targeted sequences (Servick 2016).  The U.S. Food and Drug Administration has yet to announce requirements for clinical use of the new technology.  But to help future clinicians safely repair defective, disease-causing genes, for example, researchers are exploring various means of reducing off-target effects that could harm patients in any number of ways, including through uncontrolled cellular growth and cancer.

A CRISPR-Cas9 system “licenses” a DNA sequence for cleavage through a two-stage recognition process (Bolukbasi et al. 2016). Even the most basic details are somewhat technical, of course, but very illuminating.  First, a Cas9-sgRNA complex will attach and remain attached to a DNA sequence only if an appropriate protospacer-adjacent motif (PAM) is nearby.  PAM sequences are very short, often only a few base-pairs long.  In the case of an S. pyogenes Cas9, an NGG PAM is much-preferred, but NAG and NGA PAMs are sometimes inefficiently recognized (“N” represents any nucleobase followed by two guanine, or “G” nucleobases).


Second, and only if an appropriate PAM is recognized, the sgRNA will interrogate the neighboring DNA sequence through Watson-Crick base pairing in a 3′-to-5′ direction. For an S. pyogenes Cas9, the guide sequence will measure twenty nucleotides long.  If the 3′ end of the programmed guide sequence is complementary to the DNA sequence near the PAM element, “R-loop” formation is initiated.  In zipper-like fashion, further complementarity of the DNA is assessed through extension of the R-loop.  If a complete target sequence is confirmed, allosteric activation of the Cas9 enzyme—actually, activation of Cas9’s two nuclease domains, RuvC and HNH—will result in dual cleavage and, accordingly, a complete double-stranded break in the target sequence.

Unsurprisingly, then, the specificity of a CRISPR-Cas9 system is determined in two ways. In large part, off-target effects are managed through careful design of the sgRNA.  Ideally, the guide sequence would match the target sequence perfectly, and show no homology elsewhere in the genome.  More realistically, however, at least partial homology will often occur at other genomic sites where, unfortunately, off-target cleavage could ensue.  Researchers have developed algorithms that help predict sufficient homology, but have yet to clearly and comprehensively define how closely guide and DNA sequences must harmonize before licensing occurs.  Nevertheless, nuclease activity has been observed at off-target sites displaying up to four or five nucleotide mismatches.

So, careful design of the sgRNA is critical. But one team of researchers, including Joung, recently confirmed that truncating the guide sequence can also help (Fu et al. 2014).  Shortening their guides to as few as seventeen nucleotides, instead of the usual twenty, Joung’s group was able to not only decrease nuclease activity at many off-target sites, but to preserve nearly thorough activity at the majority of intended sites as well.

Other groups have achieved similar success by inactivating one of the two nuclease domains, thus creating a “nickase” that cleaves only one strand of the target sequence (Ran et al. 2013). Here, a double-stranded break can still be achieved by joining two Cas9 nickases with two different sgRNAs targeting adjacent sites on opposing DNA strands.  Importantly, the obligatory use of two active nickases decreases the likelihood of off-target cleavage.

Perhaps the latest and most significant progress in this area, however, has been achieved through modification of the unaltered, or “wild-type,” Cas9 nuclease. Last December, for example, synthetic biologist Feng Zhang at the Broad Institute of MIT and Harvard University announced that he and his colleagues had engineered the Cas9 to render it less likely to act at genomic sites presenting mismatches between RNA guides and DNA targets (Slaymaker et al. 2015).  Appropriately, Zhang dubbed his new enzyme an “enhanced specificity” S. pyogenes Cas9, or eSpCas9 for short.

Feng Zhang

Feng Zhang

Knowing that negatively charged DNA binds to a positively charged groove in the Cas9 enzyme, Zhang’s team predicted that by replacing only a few among the 1400 or so positively charged amino acids with neutral equivalents they could temper the wild-type Cas9’s enthusiasm for binding to and cutting off-target sites. They created and tested several new versions of enzyme that reportedly reduced unintended activity at least tenfold, while maintaining robust on-target cleavage.

Earlier this year, however, Joung and colleagues claimed to have bested Zhang’s results by bringing “off-target-effects to levels where we can no longer detect them, even with the most sensitive methods” (McGreevey 2016). Like Zhang, Joung focused on points of interaction between Cas9 and DNA sequences.  His team created fifteen new enzyme variants by replacing up to four long amino acid side-chains that bind to DNA with shorter chains that do not (Kleinstiver et al. 2016).

Joung then tested each of his Cas9 variants in human cells, and found that one three-substitution and one four-substitution version rejected mismatched sites while maintaining full on-target activity. The latter variant, subsequently named SpCas9-HF1—“HF” denoting “high-fidelity,” induced targeted activity as reliably as a wild-type Cas9 when deployed with eighty-five percent of the thirty-seven different guide RNAs tested.  Similarly, SpCas9-HF1 generated no detectable off-target mutations with six of seven guide RNAs (and only one mutation with the seventh) compared to twenty-five such effects produced by the wild-type Cas9.

Keith Joung

Keith Joung

Joung’s group also tested their hi-fi creations at less typical genomic locations that are particularly difficult to control for off-target effects due to the inclusion of repeat sequences. But even there, his supplemental variants, since designated HF2, HF3, and HF4, appeared to eliminate off-target activity that tended to persist following use of the HF1 version.

It’s too early to judge which of these innovations will prove most valuable or, in fact, whether all of them will soon be superseded by modifications or entirely different systems yet to be introduced. But much progress has already been made and, importantly, at this point, many of the foregoing strategies and designs can be used in concert to bring us closer yet to the day when CRISPR gene-editing becomes a clinical convention.

Breaking Barriers.

This is now the most powerful system we have in biology. Any biological process we care about now, we can get the comprehensive set of genes that underlie that process. That was just not possible before.—biochemist David Sabatini (Yong 2015).

CRISPR-Cas9, of course, is only one among many prokaryotic CRISPR systems that could, at some point, prove useful for any number of human purposes. Use of Cas9 variations, however, has already resulted in successes far too numerous to review liberally here.  Even so, two recent applications in particular reveal the extraordinary, yet strikingly simple, means by which researchers have achieved previously unattainable outcomes.

In the first, three different teams confronted Duchenne muscular dystrophy (DMD), a terrifying disease that affects about one in every 3500 boys in the U.S. alone (Long et al. 2015, Nelson et al. 2015, and Tabebordbar et al. 2015). DMD typically stems from defects in a gene containing seventy-nine protein-coding exons.  If even a single exon suffers a debilitating mutation, the gene can be rendered incapable of producing dystrophin, a vital protein that protects muscle fibers.  Absent sufficient dystrophin, both skeletal and heart muscle will deteriorate.  Patients usually end up confined to wheelchairs and dead before the age of thirty.


Traditional gene therapy, stem cell treatments, and drugs have proven mostly ineffective against DMD. Scientists have corrected diseased cells in vitro, or in a single organ—the liver.  But treating muscle cells throughout the body, including the heart, is a far more daunting task, because they can’t all be removed, treated in isolation, and then replaced.  And given current ethical concerns, most researchers are prohibited from even considering the possibility of editing human embryos for clinical purposes.

As such, researchers here decided to employ CRISPR-Cas9 technology to excise faulty dystrophin gene exons in both adult and neonatal mice by delivering it directly into their muscles and bloodstreams using non-pathogenic adeno-associated viruses. AAVs, however, are too small to accommodate the relatively large S. pyogenes Cas9, so each team opted instead to deploy a more petite Cas9 enzyme found in Staphylococcus aureus.

Neither group’s interventions resulted in complete cures. But dystrophin production and muscle strength was restored, and little evidence of off-target effects was observed, in treated mice.  One lead researcher later suggested that, although clinical trials could be years away, up to eighty percent of human DMD victims could benefit from defective exon removal (Kaiser 2015).

Remarkably, each of the three teams obtained results comparable to those of the others. Perhaps most impressively, however, these experiments marked the very first instances of using CRISPR to successfully treat genetic disorders in fully-developed living mammals.

But an ever-growing population needs to protect its agricultural products too. Plant DNA viruses, for example, can cause devastating crop damage and economic crises worldwide, but especially in underdeveloped regions including sub-Saharan Africa.  More specifically, the tomato yellow leaf curl virus (tomato virus) is known to ravage a variety of tomato breeds, causing stunted growth, abnormal leaf development, and fruit death.


Like DMD, the tomato virus has proven an especially intractable problem. Despite previous efforts to control it through breeding, insecticides targeting the vector, and other engineering techniques, we currently know of no effective means of managing the virus.  Undeterred, another group of biologists decided to give CRISPR-Cas9-mediated viral interference a try (Ali et al. 2015).

In this study, the investigators chose to manipulate a species of tobacco plant, well-understood as a model organism, which is similarly vulnerable to tomato virus infection. The experiment was completed in two fairly predictable stages.  First, the group designed sgRNAs to target certain tomato virus coding and non-coding sequences and inserted them into different, harmless viruses of the tobacco rattle variety.  Second, they delivered the newly loaded rattle viruses into their tobacco plants.  After seven days, the plants were exposed to the tomato virus and, after ten more days, they were analyzed for symptoms of infection.

The group agreed that the CRISPR-Cas9 system had reliably cleaved and introduced mutations to the tomato viruses’ genomes. Fortuitously, every plant expressing the system had either abolished or significantly attenuated all symptoms of infection.  The investigators concluded further that the technique was capable of simultaneously targeting multiple DNA viruses with a lone sgRNA, and that other transformable plant species, including tomatoes, of course, would be similarly affected.

One can only guess, at this point, how certain interests might receive these and other types of genome-edited crops. Will nations eventually classify them as GMO or, alternatively, as organisms capable of developing in nature?  Will applicable regulations focus on the processes or products of modification?  Regardless, one can hardly ignore these commodities’ potential windfalls, especially for those in dire need.

Given recent innovations in specificity, for example, CRISPR-based disease research will likely continue to advance quickly toward clinical and other more practical applications. So long as it affects only non-reproductive somatic cells, such interventions should remain largely uncontroversial.  Human gametes and embryos, on the other hand, have once again inspired abundant debate and bitter division among experts.

Moralizing Over Science.

Genome editing in human embryos using current technologies could have unpredictable effects on future generations. This makes it dangerous and ethically unacceptable.—Edward Lanphier et al. (2015).

To intentionally refrain from engaging in life-saving research is to be morally responsible for the foreseeable, avoidable deaths of those who could have benefitted.—bioethicist Julian Savulescu et al. (2015).

The results of the first and, so far, last attempt to edit human embryos using CRISPR-Cas9 was published by a team of Chinese scientists on April 18 of last year (Liang et al. 2015). Led by Junjiu Huang, the group chose to experiment on donated tripronuclear zygotes—non-viable early embryos containing one egg and two sperm nuclei—neither intended nor suitable for clinical use.  Their goal was to successfully edit endogenous β-globin genes that, when mutated, can cause a fatal blood disorder known as β-thalassemia.

Junjiu Huang

Junjiu Huang

By his own admission, Huang’s outcomes were less than spectacular. Eighty-six embryos were injected with the Cas9 system and a molecular template designed to affect the insertion of new DNA.  Of the seventy-one that survived, fifty-four embryos were tested.  A mere twenty-eight were successfully spliced and, of those, only four exhibited the desired additions.  Rates of off-target mutations were much higher than expected too, and the group would likely have discovered additional unintended cuts had they examined more than the protein-coding exome, which represents less than two percent of the entire human genome.

In all fairness, however, the embryos’ abnormality might have been responsible for much of the total off-target effect. And, of course, Huang was unable to take advantage of many specificity-enhancing upgrades to the CRISPR system yet to be designed at the time of his investigations.  In any case, his team acknowledged that their results “highlight the pressing need to further improve the fidelity and specificity” of the new technology, which in their opinions remained immature and unready for clinical applications.

Nevertheless, the Chinese experiment ignited a brawl among both scientists and bioethicists over the prospect of human germline modification with the most powerful and accessible editing machinery ever conceived. Similar quarrels had accompanied the proliferation of technologies involving recombinant DNA, in vitro fertilization, gene therapy, and stem cells, for example.  But never had the need to address our capacity to reroute the evolution of societies—indeed, of the entire species—seemed so real and immediate.

Leading experts, including Baltimore and Doudna, had previously met in Napa, California, on January 24, 2015 to discuss the bioethical implications of rapidly emerging technologies. In the end, they “strongly discouraged … any attempts at germline genome modification for clinical application in humans,” urged informed discussion and transparent research, and called for a prompt global summit to recommend international policies (Baltimore et al. 2015).  A surge of impassioned literature ensued.

A small group led by Sangamo BioSciences president, Edward Lanphier, was one of the first to weigh in (Lanphier et al. 2015). Calling for a “voluntary moratorium” on all human germline research, Lanphier first expressed concerns over potential off-target effects and the genetic mosaicism that could result, for instance, if a fertilized egg began dividing before all intended corrections had occurred.  He also found it difficult to “imagine a situation in which use of human embryos would offer therapeutic benefits over existing and developing methods,” suggesting as well that pre-implantation genetic diagnosis (PGD) and in vitro fertilization (IVF) were far better options than CRISPR for parents carrying the same mutation for a genetic disease.  In any case, he continued, with so many unanswered questions, clinicians remained unable to obtain truly risk-informed consent from either parents looking to modify their germlines or from affected future generations.  Finally, Lanphier implied that even the best intentions could eventually lead societies down a “slippery slope” toward non-therapeutic genetic enhancement and so-called “designer babies.”

Edward Lanphier

Edward Lanphier

Francis Collins, evangelical Christian and director of the National Institutes of Health (which currently refuses to fund human germline research), expressed similar views regarding the sufficiency of PGD and IVF, the impossibility of informed consent, and non-therapeutic enhancement (Skerrett 2015). Additionally, Collins worries that access to the technology would be denied to the economically disadvantaged and that parents might begin to conceive of their children “more like commodities than precious gifts.”  For the director, given the “paucity of compelling cases” in favor of such research, and the significance of the ethical counterarguments, “the balance of the debate leans overwhelmingly against human germline engineering.”

On the other hand, Harvard Medical School geneticist, George Church, urges us to ignore pleas for artificially imposed bans, “encourage the innovators,” and focus more on what he deems the obvious benefits of germline research (Church 2015). Responding to Lanphier and Collins, he argues as well that, without obtaining consent, parents have long exposed future generations to mutagenic forces—through chemotherapy, residence in high-altitudes, and alcohol intake, for example.  We have also consistently chosen to enhance our offspring and future generations through mate choice, among many other things.  Church also points out that PGD during the IVF procedure is incapable of offering solutions to individuals possessing two copies of a detrimental, dominant allele, or to prospective parents who both carry two copies of a harmful, recessive allele.  Moreover, in most instances, PGD cannot be used to avoid more complex polygenic diseases, including schizophrenia.   Nor can we presume that new technology costs will always create treatment or enhancement inequities.  In fact, according to Church, the price of DNA sequencing, for example, has already plummeted more than three million fold.  Finally, germline editing is probably not irreversible, Church contends, and certainly not as error-prone at this point as many have suggested.  “Senseless” bans, he concludes, would only “put a damper on the best medical research and instead drive the practice underground to black markets and uncontrolled medical tourism.”

George Church

George Church

Taking a slightly different tack, Harvard cognitive scientist, Steven Pinker, censures bioethicists generally for getting bogged down in “red-tape, moratoria, or threats of prosecution based on nebulous but sweeping principles such as ‘dignity,’ ‘sacredness,’ or ‘social justice’” (Pinker 2015a). Imploring the bioethical community to “get out of the way” of CRISPR, Pinker reminds them that, once decried as morally unacceptable, vaccinations, transfusions, artificial insemination, organ transplants, and IVF have all proven “unexceptional boons to human well-being.”  Further, the specific harms of which moratorium proponents warn, including cancer, mutations, and birth defects, “are already ruled out by a plethora of existing regulations and norms” (Pinker 2015b).  In the end, he advises, both scientists and everyday people need and deserve a well-diversified research portfolio.  “If you ban something, the probability that people will benefit is zero.  If you don’t ban it, the probability is greater than zero.”

Such were among the arguments considered by a committee of twelve biologists, physicians, and ethicists during the December, 2015 International Summit on Human Genome Editing, organized by the U.S. National Academies of Science and Medicine, the Royal Society in London, and the Chinese Academy of Sciences. The Summit was chaired by David Baltimore.  Doudna and Charpentier, winners of the US$3 million Breakthrough Prize in Life Sciences, attended with Zhang—a now much-celebrated trio considered front runners for a Nobel Prize, though also entangled through their institutions in a CRISPR patent dispute potentially worth billions of dollars.

Doudna, Charpentier, and Zhang

Doudna, Charpentier, and Zhang

After three days of discussion, the Summit’s organizing committee issued a general statement rejecting calls for a comprehensive moratorium on germline research (NAS 2015). The members did, however, advise without exception against the use of edited embryos to establish pregnancy.  “It would be irresponsible to proceed,” they added, “with any clinical use of germline editing” until safety and efficacy issues are resolved and there exists “a broad societal consensus about the appropriateness of the proposed application.”  In conclusion, the committee called for an “ongoing forum” to harmonize the current global patchwork of relevant regulations and guidelines and to “discourage unacceptable activities.”  This forum, the members judged, should consist not only of experts and policymakers, but of “faith leaders,” “public interest advocates,” and “members of the general public” as well.

Wasting little time, the UK’s Human Fertilization and Embryology Authority approved on February 1, 2016, the first attempt to edit healthy human embryos with the CRISPR-Cas9 system.  The application was filed last September by developmental biologist, Kathy Niakan, of the Francis Crick Institute in London.  Niakan intends to use CRISPR to knock out one of four different genes in a total of 120 day-old, IVF-donated embryos to investigate the roles such genes play in early development.

Kathy Niakan

Kathy Niakan

Her research could help identify genes crucial to early human growth and cell differentiation and, thus, lead to more productive IVF cultures and more informed selection practices. It could also reveal mutations that lead to miscarriages and, one day, allow parents to correct these problems through gene therapy.  Following careful observation, Niakan intends to destroy her embryos by the time they reach the blastocyst stage on the seventh day.  Under British law, experimental embryos cannot be used to establish pregnancy.

But the human germline is not the only, or even most pressing, subject of CRISPR controversy. Some, for example, warn of the creation of dangerous pathogens and biological warfare (Greely 2016).  But many others, including Doudna, urge that we quickly address “other potentially harmful applications … in non-human systems, such as the alteration of insect DNA to ‘drive’ certain genes into a population” (Doudna 2015).

Driving DNA.

Clearly, the technology described here is not to be used lightly. Given the suffering caused by some species, neither is it obviously one to be ignored.—evolutionary geneticist Austin Burt (2003).

In broad terms, a “gene drive” can be characterized as a targeted contagion intended to spread through a population with exceptional haste. Burt pioneered the technology through his study of transposable elements—“selfish” and often parasitic DNA sequences that exist merely to propagate themselves.  Importantly, transposons can circumvent the normal Mendelian rules of inheritance dictating that any given gene has a fifty percent chance of being passed from parent to offspring.

Thirteen years ago, Burt envisioned the use of a microbial transposon-like element called a “homing endonuclease” for humanity’s benefit. When inserted into one chromosome, the endonuclease would cut the matching chromosome inherited from the other parent.  The cell would then quickly repair the cut, often using the first chromosome as a template.  As such, the assailed sequence in the second chromosome would be converted to the sequence of the selfish element.  In a newly fertilized egg, the endonuclease would likewise convert the other parent’s DNA and, eventually, drive itself into the genomes of nearly one-hundred percent of the population.



Burt believes we can use gene drives to weaken or even eradicate mosquito transmitted diseases like malaria and dengue fever. If scientists engineered just one percent of a mosquito population to carry such a drive, he calculates, about ninety-nine percent would possess it in only twenty generations.  In fact, Burt announced five years ago that he had created a homing endonuclease capable of locating and cutting a mosquito gene (Windbichler 2011).  But his elements were difficult to program for precise application.

Enter CRISPR-Cas9. As we’ve seen, Cas9 is an eager endonuclease and guide RNAs are easy to program and can be quickly synthesized.  In April of last year, biologists Valentonio Gantz and Ethan Bier revealed that they had used CRISPR-Cas9 to drive color variation into Drosophila fruit flies (Gantz and Bier 2015).  Though they labeled it a “mutagenic chain reaction” at the time, it was the first gene drive ever deployed in a multicellular organism.

Today, researchers sort potential gene drives into two major groups. Replacement drives seek only to displace natural with modified populations.  Suppression drives, by contrast, attempt to reduce or even eradicate populations.  At this point, no drives have been released into the wild.  Nevertheless, researchers have lately designed one of each type to affect mosquitos carrying the deadly human malaria parasite, Plasmodium falciparum.

The first study was led by microbiologist Anthony James, who collaborated on the project with Gantz and Bier (James et al. 2015). Focusing on the prevention of disease transmission, this group engineered Anopheles stephensi mosquitos, highly active in urban India, to carry two transgenes producing antibodies against the malaria parasite, a CRISPR-Cas9-mediated gene drive, and a marker gene.  Because the very lengthy payload rendered insertion a challenging process, James was able to isolate only two drive-bearing males among 25,000 larvae.  But when mated with wild-type females, these and subsequent transgenic males spread their anti-malaria genes at an impressive rate of 99.5 percent.  Transgenic females, on the other hand, processed the drive quite differently and passed it on at near-normal Mendelian ratios.

Despite its overall success, James doesn’t imagine that his team’s replacement drive could eliminate the malaria parasite independently. Instead, he envisions its use to reduce the risk of infection and to compliment other strategies already being employed.  Even so, because such drives would not exterminate P. falciparum or its mosquito vector, they would potentially allow the parasite to one day evolve resistance to their transgene components.


The second study’s goal was quite different. Here, molecular biologist, Tony Nolan, along with Burt and others, first identified three genes in the Anopheles gambiae mosquito, active in sub-Saharan Africa, that when mutated cause recessive infertility in females (Hammond et al. 2016).  Second, they designed a CRISPR-Cas9 gene drive to target and edit each gene.  Following insertion, they bred their transgenic mosquitos with wild-types and found that nearly all female offspring were born infertile.  In a subsequent experiment, Nolan released 600 vectors—half transgenic, half wild-type—into a cage.  After only four generations, seventy-five percent of the population carried the mutations, exactly what one would expect from an effective gene drive.

A suppression drive like Hammond’s could, in theory, eliminate a parasite’s primary vector. In such a scenario, the parasite might find another means of conveying the disease to humans—more than 800 species of mosquito inhabit Africa alone, for example.  But it might not.  The loss would also substantially alter the relevant ecosystem.  But despite other methods of controlling the disease, malaria still claims more than a half million lives every year, mostly among children under five.

Even in theory, no gene drive is a panacea. They function only in sexually reproducing species, and best in species that reproduce very rapidly.  Nor would their effects be permanent—most transgenes would prove especially vulnerable to evolutionary deselection, for example.   But neither would they turn out as problematic as some might imagine. They can be easily detected through genome sequencing, for instance, and are unlikely to spread accidentally into domesticated species.  And if scientists sought for whatever reason to reverse the effects of a previously released drive, they could probably do so with the release of a subsequent drive.

As Church and others have recently suggested, it “doesn’t really make sense to ask whether we should use gene drives. Rather, we’ll need to ask whether it’s a good idea to consider driving this particular change through this particular population”  (Esvelt et al. 2014).


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Biological Race and the Problem of Human Diversity (Cover Article).

by Kenneth W. Krause.

Kenneth W. Krause is a contributing editor and “Science Watch” columnist for the Skeptical Inquirer.  Formerly a contributing editor and books columnist for the Humanist, Kenneth contributes regularly to Skeptic as well.  He may be contacted at

Race 1

Some would see any notion of “race” recede unceremoniously into the dustbin of history, taking its ignominious place alongside the likes of phlogiston theory, Ptolemaic geocentricism, or perhaps even the Iron Curtain or Spanish Inquisition.  But race endures, in one form or another, despite its obnoxious, though apparently captivating dossier.

In 1942, anthropologist Ashley Montagu declared biological race “Man’s Most Dangerous Myth,” and, since then, most scientists have consistently agreed (Montagu 1942).  Nevertheless, to most Americans in particular, heritable race seems as obvious as the colors of their neighbors’ skins and the textures of their hair.  So too have a determined minority of researchers always found cause to dissent from the professional consensus.

Here, I recount the latest popular skirmish over the science of race and attempt to reveal a victor, if there be one.  Is biological race indeed a mere myth, as the academic majority has asked us to concede for more than seven decades?  Is it instead a scandalously inconvenient truth—something we all know exists but, for whatever reasons, prefer not to discuss in polite company?  Or is it possible that a far less familiar rendition of biological race could prove not only viable, but both scientifically and socially valuable as well?

Race Revived.

The productive questions pertain to how races came to be and the extent to which racial variation has significant consequences with respect to function in the modern world.—Vincent Sarich and Frank Miele, 2004.

I have no reason to believe that Nicholas Wade, long-time science editor and journalist, is a racist, if “racist” is to mean believing in the inherent superiority of one human race over any other.  In fact, he expressly condemns the idea.  But in the more limited and hopefully sober context of the science of race, Wade is a veritable maverick.  Indeed, his conclusions that biological human races (or subspecies, for these purposes) do exist, and conform generally to ancestral continental regions, appear remarkably more consistent with those of the general public.

In his most recent and certainly controversial book, A Troublesome Inheritance: Genes, Race and Human History, Wade immediately acknowledges that the vast majority of both anthropologists and geneticists deny the existence of biological race (Wade 2014).  Indeed, “race is a recent human invention,” according to the American Anthropological Association (AAA 2008), and a mere “social construct,” per the American Sociological Association (ASA 2003).  First to decode the human genome, Craig Venter was also quick to announce during his White House visit in 2000 that “the concept of race has no genetic or scientific basis.”

But academics especially are resistant to biological race, or the idea that “human evolution is recent, copious, and regional,” Wade contends, because they fear for their careers in left-leaning political atmospheres and because they tend to be “obsessed with intelligence” and paralyzed by the “unlikely” possibility that genetics might one day demonstrate the intellectual superiority of one major race over others.

According to Wade, “social scientists often write as if they believe that culture explains everything and race [indeed, biology] explains nothing, and that all cultures are of equal value.”  But “the emerging truth,” he insists, “is more complicated.”  Although the author sees individuals as fundamentally similar, “their societies differ greatly in their structure, institutions and their achievements.”  Indeed, “contrary to the central belief of multiculturalists, Western culture has achieved far more” than others “because Europeans, probably for reasons of both evolution and history, have been able to create open and innovative societies, starkly different from the default human arrangements of tribalism or autocracy.”

Race 6

Wade admits that much of his argument is speculative and has yet to be confirmed by hard, genetic evidence.  Nevertheless, he argues, “even a small shift in [genetically-based] social behavior can generate a very different kind of society,” perhaps one where trust and cooperation can extend beyond kin or the tribe—thus facilitating trade, for example, or one emphasizing punishment for nonconformity—thus advancing rule-orientation and isolationism, for instance.  “[I]t is reasonable to assume,” the author vies, “that if traits like skin color have evolved in a population, the same may be true of its social behavior.”

But what profound environmental conditions could possibly have selected for more progressive behavioral adaptations in some but not all populations?  As the climate warmed following the Pleistocene Ice Age, Wade reminds, the agricultural revolution erupted around 10,000 years ago among settlements in the Near East and China.  Increased food production led to population explosions, which in turn spurred social stratification, wealth disparities, and more frequent warfare.  “Human social behavior,” Wade says, “had to adapt to a succession of makeovers as settled tribes developed into chiefdoms, chiefdoms into archaic states and states into empires.”

Meanwhile, other societies transformed far less dramatically.  “For lack of good soils, favorable climate, navigable rivers and population pressures,” Wade observes, “Africa south of the Sahara remained largely tribal throughout the historical period, as did Australia, Polynesia and the circumpolar regions.”

Citing economist Gregory Clark, Wade then postulates that, during the period between 1200 and 1800 CE—twenty-four generations and “plenty of time for a significant change in social behavior if the pressure of natural selection were sufficiently intense,”—the English in particular evolved a greater tendency toward “bourgeoisification” and at least four traits—nonviolence, literacy, thrift, and patience—thus enabling them to escape the so-called “Malthusian trap,” in which agrarian societies never quite learn to produce more than their expanding numbers can consume, and, finally, to lead the world into the Industrial Revolution.

In other words, according to this author, modern industrialized societies have emerged only as a result of two evolved sets of behaviors—initially, those that favor broader trust and contribute to the breakdown of tribalism, and, subsequently, those that favor discipline and delayed gratification and lead to increased productivity and wealth.  On the other hand, says Wade, Sub-Saharan Africans, for example, though well-adapted to their unique environmental circumstances, generally never evolved traits necessary to move beyond tribalism.  Only an evolutionary explanation for this disparity, he concludes, can reveal, for instance, why foreign aid to non-modern societies frequently fails and why Western institutions, including democracy and free markets, cannot be readily transferred to (or forced upon) yet pre-industrial cultures.

So how many races have evolved in Wade’s estimation?  Three major races—Caucasian, East Asian, and African—resulted from an early migration out of Africa some 50,000 years ago, followed by a division between European and Asian populations shortly thereafter.  Quoting statistical geneticist, Neil Risch, however, Wade adds Pacific Islanders and Native Americans to the list because “population genetic studies have recapitulated the classical definition of races based on continental ancestry” (Risch 2002).

To those who would object that there can be no biological race when so many thousands of people fail to fit neatly into any discreet racial category, Wade responds, “[T]o say there are no precise boundaries between races is like saying there are no square circles.”  Races, he adds, are merely “way stations” on the evolutionary road toward speciation.  Different variations of a species can arise where different populations face different selective challenges, and humans have no special exemption from this process.  However, the forces of differentiation can reverse course when, as now, races intermingle due to increased migration, travel, and intermarriage.

Race Rejected.

It is only tradition and shortsightedness that leads us to think there are multiple distinct oceans.—Guy P. Harrison, 2010.

So, if we inherit from our parents traits typically associated with race, including skin, hair, and eye color, why do most scientists insist that race is more social construct than biological reality?  Are they suffering from an acute case of political correctness, as Wade suggests, or perhaps a misplaced paternalistic desire to deceive the irresponsible and short-sighted masses for the greater good of humanity?  More ignoble things have happened, of course, even within scientific communities. But according to geneticist Daniel J. Fairbanks, the denial of biological race is all about the evidence.

In his new book, Everyone is African: How Science Explodes the Myth of Race, Fairbanks points out that, although large-scale analyses of human DNA have recently unleashed a deluge of detailed genetic information, such analyses have so far failed to reveal discrete genetic boundaries along traditional lines of racial classification (Fairbanks 2015).  “What they do reveal,” he argues, “are complex and fascinating ancestral backgrounds that mirror known historical immigration, both ancient and modern.”


In 1972, Harvard geneticist Richard Lewontin analyzed seventeen different genes among seven groups classified by geographic origin.  He famously discovered that subjects within racial groups varied more among themselves than their overall group varied from other groups, and concluded that there exists virtually no genetic or taxonomic significance to racial classifications (Lewontin 1972).  But Lewontin’s word on the subject was by no means the last. Later characterizing his conclusion as “Lewontin’s Fallacy,” for example, Cambridge geneticist A.W.F. Edwards reminded us how easy it is to predict race simply by inspecting people’s genes (Edwards 2003).

So who was right?  Both of them were, according to geneticist Lynn Jorde and anthropologist Stephen Wooding.  Summarizing several large-scale studies on the topic in 2004, they confirmed Lewontin’s finding that about 85-90% of all human genetic variation exists within continental groups, while only 10-15% between them (Jorde and Wooding 2004).  Even so, as Edwards had insisted, they were also able to assign all native European, East Asian, and sub-Saharan African subjects to their continent of origin using DNA alone.  In the end, however, Jorde and Wooding showed that geographically intermediate populations—South Indians, for example—did not fit neatly into commonly conceived racial categories.  “Ancestry,” they concluded, was “a more subtle and complex description” of one’s genetic makeup than “race.”

Fairbanks concurs.  Humans have been highly mobile for thousands of years, he notes.  As a result, our biological variation “is complex, overlapping, and more continuous than discreet.”  When one analyzes DNA from a geographically broad and truly representative sample, the author surmises, “the notion of discrete racial boundaries disappears.”

Nor are the genetic signatures of typically conceived racial traits always consistent between populations native to different geographic regions.  Consider skin color, for example.  We know, of course, that the first Homo sapiens inherited dark skin previously evolved in Africa to protect against sun exposure and folate degradation, which negatively affects fetal development.  Even today, the ancestral variant of the MC1R gene, conferring high skin pigmentation, is carried uniformly among native Africans.

Race 2

But around 30,000 years ago, Fairbanks instructs, long after our species had first ventured out of Africa into the Caucasus region, a new variant appeared.  KITLG evolved in this population prior to the European-Asian split to reduce pigmentation and facilitate vitamin D absorption in regions of diminished sunlight.  Some 15,000 years later, however, another variant, SLC24A5, evolved by selective sweep as one group migrated westward into Europe.  Extremely rare in other native populations, nearly 100% of modern native Europeans carry this variant.  On the other hand, as their assorted skin tones demonstrate, African and Caribbean Americans carry either two copies of an ancestral variant, two copies of the SLC24A5 variant, or one of each.  Asians, by contrast, developed their own pigment-reducing variants—of the OCA2 gene, for example—via convergent evolution, whereby similar phenotypic traits result independently among different populations due to similar environmental pressures.

So how can biology support the traditional, or “folk,” notion of race when the genetic signatures of that notion’s most relied upon trait—that is, skin color—are so diverse among populations sharing the same or similar degree of skin pigmentation?  Fairbanks judges the idea utterly bankrupt “in light of the obvious fact that actual variation for skin color in humans does not fall into discrete classes,” but rather “ranges from intense to little pigmentation in continuously varying gradations.”

To Wade, Fairbanks offers the following reply: “Traditional racial classifications constitute an oversimplified way to represent the distribution of genetic variation among the people of the world. Mutations have been creating new DNA variants throughout human history, and the notion that a small proportion of them define human races fails to recognize the complex nature of their distribution.”

Race 8

A Severe Response.

I use the term scientific racism to refer to scientists who continue to believe that race is a biological reality.—Robert Wald Sussman, 2014.

Since neither author disputes the absence of completely discreet racial categories, one could argue that part of the battle is really one over mere semantics, if not politics. Regardless, critical aspects of Wade’s analysis were quickly and sharply criticized by several well-respected researchers.

Former president of the AAA and co-drafter of its statement on race, Alan Goodman, for example, argues that Wade’s “speculations follow from misunderstandings about most everything, including the idea of race, evolution and gene action, culture and institutions, and most fundamentally, the scientific process” (Goodman 2014). Indeed, he compares Wade’s book to the most maligned texts on race ever published, including Madison Grant’s 1916 The Passing of the Great Race, Arthur Jensen’s 1969 paper proposing racial intelligence differences, and Herrnstein’s and Murray’s 1994 The Bell Curve.

But Wade’s “biggest error,” according to Goodman, “is his inability to separate the data on human variation from race.” He mistakenly assumes, in other words, “that all he sees is due to genes,” and that culture means little to nothing. A “mix of mysticism and sociobiology,” he continues, Wade’s simplistic view of human culture ignores the archeological and historical fact that cultures are “open systems” that constantly change and interact. And although biological human variation can sometimes fall into geographic patterns, Goodman emphasizes, our centuries-long attempt to force all such variation into racial categories has failed miserably.

Characterizing Wade’s analysis similarly as a “spectacular failure of logic,” population geneticist Jennifer Raff takes special issue with the author’s attempt to cluster human genetic variation into five or, really, any given number of races (Raff 2014). To do so, Wade relied in part on a 2002 study featuring a program called Structure, which is used to group people across the globe based on genetic similarities (Rosenberg 2002). And, indeed, when Rosenberg et al. asked Structure to bunch genetic data into five major groups, it produced clusters conforming to the continents.

But, as Raff observes, the program was capable of dividing the data into any number of clusters, up to twenty in this case, depending on the researchers’ pre-specified desire. When asked for six groups, for example, Structure provided an additional “major” cluster, the Kalash of northwestern Pakistan—which Wade arbitrarily, according to Raff, rejected as a racial category. In the end, she concludes, Wade seems to prefer the number five “simply because it matches his pre-conceived notions of what race should be.”

Interestingly, when Rosenberg et al. subsequently expanded their dataset to include additional genetic markers for the same population samples, Structure simply rejected the Kalesh and decided instead that one of Wade’s five human races, the Native Americans, should be split into two clusters (Rosenberg 2005). In any event, Rosenberg et al. expressly warned in their second paper that Structure’s results “should not be taken as evidence of [their] support of any particular concept of ‘biological race.’”

Structure was able to generate discrete clusters from a very limited quantity of genetic variation, adds population geneticist Jeremy Yoder, because its results reflect what his colleagues refer to as isolation-by-distance, or the fact that populations separated by sufficient geographic expanses will display genetic distinctions even if intimately connected through migration and interbreeding (Yoder 2014). In reality, however, human genetic variation is clinal, or gradual in transition between such populations. In simpler terms, people living closer together tend to be more closely related than those living farther apart.

In his review, biological anthropologist Greg Laden admits that human races might have existed in the past and could emerge at some point in the future (Laden 2014). He also concedes that “genes undoubtedly do underlie human behavior in countless ways.” Nevertheless, he argues, Wade’s “fashionable” hypothesis proposing the genetic underpinnings of racially-based behaviors remains groundless. “There is simply not an accepted list of alleles,” Laden reminds, “that account for behavioral variation.”

Chimpanzees, by contrast, can be divided into genetically-based subspecies (or races). Their genetic diversity has proven much greater than ours, and they demonstrate considerable cultural variation as well. Even so, Laden points out, scientists have so far been unable to sort cultural variation among chimps according to their subspecies. So if biologically-based races cannot explain cultural differences among chimpanzees, despite their superior genetic diversity as a species, why would anyone presume the opposite of humans?

Race 7

None of which is to imply that every review of Wade has been entirely negative. Conservative journalist Anthony Daniels (a.k.a. Theodore Dalrymple), for example, praises the author lavishly as a “courageous man … who dares raise his head above the intellectual parapet” (Daniels 2014). While judging Wade’s arguments mostly unconvincing, he nevertheless defends his right to publish them: “That the concept of race has been used to justify the most hideous of crimes should no more inhibit us from examining it dispassionately … than the fact that economic egalitarianism has been used to justify crimes just as hideous …”

Similarly, political scientist and co-author of The Bell Curve, Charles Murray warned readers of the social science “orthodoxy’s” then-impending attempt to “not just refute” Wade’s analysis, “but to discredit it utterly—to make people embarrassed to be seen purchasing it in public” (Murray 2014). “It is unhelpful,” Murray predicts, “for social scientists and the media to continue to proclaim that ‘race is a social construct’” when “the problem facing us down the road is the increasing rate at which the technical literature reports new links between specific genes and specific traits.” Although “we don’t yet know what the genetically significant racial differences will turn out to be,” Murray contends, “we have to expect that they will be many.”

Perhaps; perhaps not. But race is clearly problematic from a biological perspective—at least as Wade and many before him have imagined it. Humans do not sort neatly into separate genetic categories, or into a handful of continentally-based groups. Nor have we discovered sufficient evidence to suggest that human behaviors match to known patterns of genetic diversity. Nonetheless, because no “is” ever implies an “ought,” the cultural past should never define, let alone restrain, the scientific present.

Characterizing Biological Diversity.

Instead of wasting our time “refuting” straw-man positions dredged from a distant past or from fiction, we should deal with the strongest contemporary attempts to rehabilitate race that are scientifically respectable and genetically informed.—Neven Sesardic, 2010.

To this somewhat belated point, I have avoided the task of defining “biological race,” in large measure because no single definition has achieved widespread acceptance. In any event, preeminent evolutionary biologist, Ernst Mayr, once described “geographic race” generally as “an aggregate of phenotypically similar populations of a species inhabiting a geographic subdivision of the range of that species and differing taxonomically from other populations of that species” (Mayr 2002). A “human race,” he added, “consists of the descendants of a once-isolated geographic population primarily adapted for the environmental conditions of their original home country.”

Sounds much like Wade, so far. But unlike Wade, Mayr firmly rejected any typological, essentialist, or folk approach to human race denying profuse variability and mistaking non-biological attributes—especially those implicating personality and behavior—for racial traits. Accepting culture’s profound sway, Mayr warned that it is “generally unwise to assume that every apparent difference … has a biological cause.” Nonetheless, he concluded, recognizing human races “is only recognizing a biological fact”:

Geographic groups of humans, what biologists call races, tend to differ from each other in mean differences and sometimes even in specific single genes. But when it comes to the capacities that are required for the optimal functioning of our society, I am sure that any racial group can be matched by that of some individual in another racial group. This is what population analysis reveals.

So how might one rescue biological race from the present-day miasma of popular imparsimony and professional denialism, perhaps even to the advancement of science and benefit of society? Evolutionary biologist and professor of science philosophy, Massimo Pigliucci, thinks he has an answer.

Race 3

More than a decade ago, he and colleague Jonathan Kaplan proposed that “the best way of making sense of systematic variation within the human species is likely to rely on the ecotypic conception of biological races” (Pigliucci and Kaplan 2003). Ecotypes, they specify, are “functional-ecological entities” genetically adapted to certain environments and distinguished from one another based on “many or a very few genetic differences.” Consistent with clinal variation, ecotypes are not always phylogenetically distinct, and gene flow between them is common. Thus, a single population might consist of many overlapping ecotypes.

All of which is far more descriptive of human evolution than even the otherwise agreeable notion of “ancestry,” for example. For Pigliucci and Kaplan, the question of human biological race turns not on whether there exists significant between-population variation overall, as Lewontin, for example, suggested, but rather on “whether there is [any] variation in genes associated with significant adaptive differences between populations.” As such, if we accept an ecotypic description of race, “much of the evidence used to suggest that there are no biologically significant human races is, in fact, irrelevant.”

On the other hand, as Pigliucci observed more recently, the ecotypic model implies the failure of folk race as well. First, “the same folk ‘race’ may have evolved independently several times,” as explained above in the context of skin color, “and be characterized by different genetic makeups” (Pigliucci 2013). Second, ecotypes are “only superficially different from each other because they are usually selected for only a relatively small number of traits that are advantageous in certain environments.” In other words, the popular notion of the “black race,” for example, centers on a scientifically incoherent unit—one “defined by a mishmash of small and superficial set of biological traits … and a convoluted cultural history” (Pigliucci 2014).

So, while the essentialist and folk concepts of human race can claim “no support in biology,” Pigliucci concludes, scientists “should not fall into the trap of claiming that there is no systematic variation within human populations of interest to biology.” Consider, for a moment, the context of competitive sports. While the common notion that blacks are better runners than whites is demonstrably false, some evidence does suggest that certain West Africans have a genetic edge as sprinters, and that certain East and North Africans possess an innate advantage as long-distance runners (Harrison 2010). As the ecotypic perspective predicts, the most meaningful biological human races are likely far smaller and more numerous than their baseless essentialist and folk counterparts (Pigliucci and Kaplan 2003).

So, given the concept’s exceptionally sordid history, why not abandon every notion of human race, including the ecotypic version? Indeed, we might be wise to avoid the term “race” altogether, as Pigliucci and Kaplan acknowledge. But if a pattern of genetic variation is scientifically coherent and meaningful, it will likely prove valuable as well. Further study of ecotypes “could yield insights into our recent evolution,” the authors urge, “and perhaps shed increased light onto the history of migrations and gene flow.” By contrast, both the failure to replace the folk concept of race and the continued denial of meaningful patterns of human genetic variation have “hampered research into these areas, a situation from which neither biology nor social policy surely benefit.”


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Fairbanks, D.J. 2015. Everyone is African: how science explodes the myth of race. Amherst, NY: Prometheus Books.

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Harrison, G.P. 2010. Race and reality: what everyone should know about our biological diversity. Amherst, NY: Prometheus Books.

Jorde, L.B. and S.P. Wooding. 2004. Genetic variation, classification and ‘race.’ Nature Genetics 36(11):528-533.

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Mayr, E. 2002. The biology of race and the concept of equality. Daedalus 131(1):89-94.

Montagu, A. 1942. Man’s most dangerous myth: the fallacy of race. NY: Columbia University Press.

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Pigliucci, M. 2013. What are we to make of the concept of race? Thoughts of a philosopher-scientist. Studies in History and Philosophy of Biological and Biomedical Sciences. 44:272-277.

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Rosenberg, N., M. Saurabh, S. Ramachandran, et al. 2005. Clines, clusters, and the effect of study design on the inference of human population structure. PLOS Genetics 1(6):e70.

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Sussman, R.W. 2014. The myth of race: the troubling persistence of an unscientific idea. Cambridge, MA: Harvard University Press.

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Nature, Nurture, and the Folly of “Holistic Interactionism.”

[Notable New Media]

by Kenneth W. Krause.

Kenneth W. Krause is a contributing editor and “Science Watch” columnist for the Skeptical Inquirer.  Formerly a contributing editor and books columnist for the Humanist, Kenneth contributes regularly to Skeptic as well.  He may be contacted at

Most contemporary scientists, according to Harvard University experimental psychologist, Steven Pinker, have abandoned both the nineteenth-century belief in biology as destiny and the twentieth-century doctrine that the human mind begins as a “blank slate.”  In his new anthology, Language, Cognition, and Human Nature: Selected Articles (Oxford 2015), Pinker first reminds us of the now-defunct blank slate’s political and moral appeal:  “If nothing in the mind is innate,” he chides, “then differences among races, sexes, and classes can never be innate, making the blank slate the ultimate safeguard against racism, sexism, and class prejudice.”


Even so, certain angry ideologues, for example, continue to wallow in blank slate dogma.  Gender differences in STEM professions, for example, are often attributed entirely to prejudice and hidden barriers.  The mere possibility that women, on average, are less interested than men in people-free pursuits remains oddly “unspeakable,” says Pinker (but see a recent exception here).  The point, he clarifies, is not that we know for certain that evolution and genetics are relevant to explaining so-called “underrepresentation” in high-end science and math, but that “the mere possibility is often treated as an unmentionable taboo, rather than as a testable hypothesis.”

A similar exception to the general rule centers around parenting and the behavior of children.  It may be true that parents who spank raise more violent children, and that more conversant parents produce children with better language skills.  But why does “virtually everyone” conclude from such facts that the parent’s behavior causes that of the child?  “The possibility that the correlations may rise from shared genes is usually not even mentioned, let alone tested,” says Pinker.

Equally untenable for the author is the now-popular academic doctrine he dubs “holistic interactionism” (HI).  Carrying a “veneer of moderation [and] conceptual sophistication,” says Pinker, HI is based on a few “unexceptional points,” including the facts that nature and nurture are not mutually exclusive and that genes cannot cause behavior directly.  But we should confront this doctrine with heightened scrutiny, according to Pinker, because “no matter how complex the interaction is, it can be understood only by identifying the components and how they interact.”  HI “can stand in the way of such an understanding,” he warns, “by dismissing any attempt to disentangle heredity and environment as uncouth.”

HI mistakenly assumes, for example, that hereditary cannot constrain behavior because genes depend critically on the environment.  “To begin with,” says Pinker, “it is simply not true that any gene can have any effect in some environment, with the implication that we can always design an environment to produce whatever outcome we value.”  And even if some extreme “gene-reversing” environment can be imagined, it simply doesn’t follow that “the ordinary range of environments will [even] modulate that trait, [or that] the environment can explain the nature of the trait.”  The mere existence of environmental mitigations, in other words, does not render the effects of genes inconsequential.  To the contrary, Pinker insists, “genes specify what kinds of environmental manipulations will have what kinds of effects and with what costs.”

Although the postmodernists and social constructionists who tend to dominate humanities departments in American Universities especially, continue to tout HI as a supposedly nuanced means of comprehending the nature-nurture debate, it is in truth little more than a pseudo-intellectual “dodge,” Pinker concludes: a convenient means to “evade fundamental scientific problems because of their moral, emotional, and political baggage.”

Among intellectually honest, truly curious, and consistently rational thinkers (a diminutive demographic indeed), Pinker’s reputation is and has long stood as something perhaps just short of heroic, in no small part due to his defense of politically incorrect but nonetheless scientifically viable hypotheses.  What a shame it is that only academics of similar status (and tenure) can safely rise and demand the freedom required to mount such defenses.  And what a tragedy that so few in such privileged company actually do.


Book Review: Phillip E. Hammond, David W. Machacek, and Eric Michael Mazur, Religion on Trial: How Supreme Court Trends Threaten Freedom of Conscience in America (AltaMira 2004) 177 pp.

by Kenneth W. Krause.

Kenneth W. Krause is a contributing editor and “Science Watch” columnist for the Skeptical Inquirer.  Formerly a contributing editor and books columnist for the Humanist, Kenneth contributes regularly to Skeptic as well.  He may be contacted at

“The First Amendment,” scholars of religion Hammond, Machacek, and Mazur argue, “only reaffirmed what [James] Madison understood already to be the case, which was that the federal government was powerless over conscience . . ..”  The founders, they insist, conceived of “freedom of conscience” not simply as a restatement of religious liberty, but rather as a much more comprehensive franchise—“the freedom of the individual to decide for him- or herself questions of morality, truth, and beauty,” irrespective of the individual’s religious proclivities or lack thereof.  In broad terms, the authors rest their claim upon two distinct foundations, the first solid and enduring, the second flimsy and ephemeral.

History, they profess, clearly demonstrates the founders’ inclusive aspirations.  Madison understood that ratification of the Bill of Rights would not conclude America’s struggle to define freedom of conscience.  His fondest hope for the First Amendment, in fact, was that it “would set in motion a process of expanding liberty.”  Indeed, mounting freedoms would seem vital to the survival of any emerging republic, or, otherwise stated, crucial to political and philosophical commerce within any sincerely free marketplace of ideas.

But the Supreme Court, say the authors, quickly subverted whatever hopes the earliest generations of forward-thinking Americans may have entertained with respect to the founders’ original intent.  “[N]ineteenth-century jurists,” in fact, were the culprits “who laid the foundation for the argument that conscience necessarily meant religion (and that religion meant Christianity).”  In his 1833 Commentaries on the Constitution, for example, Justice Joseph Story wrote that the First Amendment was designed “not to countenance . . . infidelity, . . .” but rather “to exclude all rivalry among Christian sects, and to prevent any national ecclesiastical establishment . . ..”

Consistent with his Protestant faith, Story distinguished religious conduct from religious conviction, sacrificing the former to the discretion of state governments and awarding the latter to the exclusive prerogative of individuals.  More than the framers’ Enlightenment values, this non-preferential “Republican Protestantism” formed the basis of the Court’s initially stubborn reluctance to involve itself in religious issues.

As immigration fueled religious diversity, however, American Protestantism gradually and grudgingly ceded its privileged legal status.  By the end of the century, for example, local court decisions involving blasphemy prosecutions, Sunday closing laws, and church property disputes reflected decidedly less bias toward an exclusively Protestant worldview.  Somewhat ironically, as Catholics and Quakers threatened to take advantage of government funding, resentful Protestants began to clamor for the separation of church and state.

Even so, the authors point out, “it would take a revolution” on the Supreme Court “to bring about a fuller understanding of the individual rights of conscience . . . articulated by the Framers over one hundred years earlier.”  Such transformation commenced following Roosevelt’s reelection in 1936 and, given the prevailing force of Republican Protestantism, relied heavily upon certain religious organizations’ assertion of their rights to free speech and thought rather than religious liberty.  “[W]hen the organization based its arguments either on a combination of free speech and religious liberty rights, or on free speech alone, [it] was much more likely to triumph.”

The Jehovah’s Witnesses, of course, played a key role in this revolution, facilitating post-New Deal America’s penchant toward the conceptual affiliation between free speech, free thought, free conscience, and religious liberty.  By the end of the Warren and Burger eras, the Court had recognized that a tribunal’s regulation of minority religious behavior and its corresponding deference to majority actions could be seen as a patently unconstitutional “casting of judges as ‘theologians’ . . ..”  Forced to abandon Republican Protestantism’s belief-action distinction, the authors argue, the Court expanded the right to free exercise laterally to accommodate conscience or conviction, regardless of whether it was expressed in religious terms.

According to Justice Harry Blackmun, writing for the Court in 1989, “Perhaps in the early days of the Republic [the First Amendment was] understood to protect only the diversity within Christianity, but today [it is] recognized as guaranteeing religious liberty and equality to the infidel, the atheist, or the adherent of a non-Christian faith . . ..”  Implicit in Blackmun’s analysis, the authors maintain, was the Justice’s understanding that official endorsement also amounts to a violation of free exercise, “because [endorsement] communicates to non-Christians that their religions are not endorsed” and to dissenting Christians that their religion ought to be imagined and practiced in a particular way.

In the authors’ estimation, Blackmun’s expanded vision of what is protected under the Free Exercise Clause and proscribed under the Establishment Clause “represents the gradual unfolding of the true meaning and purpose of the First Amendment.”  Conversely, the approach of contemporary non-preferentialists, including the “regressive bloc” of the Supreme Court which, at the time of publication, consisted of Justices Rehnquist, Thomas, and Scalia, is marked by an ahistorical analysis and the rejection of the co-dependent nature of the First Amendment’s two religion clauses.

During recent years, of course, the “enlightened” trend has decelerated and, arguably, reversed itself.  Considering each Justice’s record of liberality with respect to free exercise and conservatism in relation to establishment, the authors conclude—not unpredictably—that William O. Douglass was the most and that Clarence Thomas has been the least progressive Supreme Court jurist in history.  Similarly, the Warren Court represented the apogee of enlightenment and the 2004 Rehnquist Court its nadir.

The Court is now inclined, say the authors, toward the abandonment of strict scrutiny in free-exercise cases and the abolishment of the Lemon test in establishment cases.  The trend, in other words, entails the continued and perhaps expanded constitutional validation of ostensibly “neutral” legislation on the one hand, and “devolution,” or the philosophical infatuation with local as opposed to federal power, on the other.  The 2004 Supreme Court, they conclude, “would gladly submit the rights to ‘life, liberty, and property’ to the popular vote—precisely what the Fifth and Fourteenth Amendments [were] designed to prevent.”

The authors’ historical argument, once again, is credible and well founded.  Clearly, the First Amendment’s primary authors anticipated both the expansion of religious liberty and a much more capacious separation of religion and government than was achievable in eighteenth-century America.  The authors’ second argument, however, that the necessity of free exercise and separation is and has always been based on every human’s “natural rights,” is seriously flawed.

Although the authors claim to conceive of natural rights in individualistic Lockeian rather than religious Thomistic terms, their conclusion is no less irrational for the distinction.  “The U.S. Constitution,” they contend, “is fundamentally a ‘sacred,’ not a ‘secular’ document.”  “Freedom of conscience,” they add, “does not derive from our particular system of government; it is, rather, an aspect of ‘personality.’”

Such sentiments, if accepted as true, might console Americans who, regardless of politics or ideology, either favor a more expansive interpretation of the Free Exercise Clause or favor greater separation of religion and government and wish to base their hopes in that respect upon the attendant claim that such separation is essential to generally expanded religious liberty.

But wanting a proposition to be true does not affect that proposition’s veracity, no matter how prevailing, passionate, or appreciable the desire.  So, while the authors are correct that “to be inviolate,” freedom of conscience must be held “free from the interference of the state,” both reason and history are clear that there exists no factual basis for the concept of natural rights and that freedom of conscience, in fact, can be violated by the government and, all too often, egregiously so.

Such is precisely the reason why our civil rights remain fragile, after all, and why every fresh generation of Americans must fight for their civil rights’ continued existence.  In a society built upon Enlightenment ideals, no claim is exempt from rational inquiry and, accordingly, no idea or opinion can ever be held sacred.  Absent a factual basis, the patronizing and pretentious concept of natural rights is “simply rhetorical nonsense,” as Jeremy Bentham once suggested—“nonsense upon stilts.”

Book Review: Francis Fukuyama, America at the Crossroads: Democracy, Power, and the Neoconservative Legacy (Yale UP 2006). 226 pp.

by Kenneth W. Krause.

Kenneth W. Krause is a contributing editor and “Science Watch” columnist for the Skeptical Inquirer.  Formerly a contributing editor and books columnist for the Humanist, Kenneth contributes regularly to Skeptic as well.  He may be contacted at

Johns Hopkins international studies guru, Francis Fukuyama, has jumped an embattled neoconservative ship.  In America at the Crossroads, the author proposes a more “realistic Wilsonianism,” rejecting both the isolationist tendencies of Jacksonian nationalism and the stubborn disregard of other nations’ internal affairs characteristic of Kissingeresque realism.  Like many contemporary neocons, Fukuyama defends American military power as an occasionally appropriate means to a moral end, but, like liberal internationalists, he emphasizes soft power, multilateralism, and the use of global institutions.

Nevertheless, neither preventive war, international social programs, nor the Bush Doctrine are compatible with Fukuyama’s eclectic worldview.  Consistent with the teachings of Leo Strauss, Fukuyama defines “regime” broadly to encompass not only a society’s political authority but its cultural underpinnings as well.  Even so, the author recommends that “certain political problems can be solved only through regime change.”  But such intercessions, he warns, are not easily accomplished, certainly much more difficult than the Bush administration appeared to believe before its invasion of Iraq.

Even a superficial understanding of Islam’s history and doctrine would have dissuaded the Bush administration from attempting to impose Enlightenment values upon Iraq in such brisk and brutish order.  As Benjamin Barber recently observed, no government can export democracy because no society can import civil rights.  Such institutions must evolve slowly and from deep inside a culture’s gut.

“This makes an exclusively military response to the challenge inappropriate,” Fukuyama concurs.  America’s record with regard to international democratization has been spotty at best.  Prior to the second Iraq war, Bush administration officials peddled Germany and Japan as obvious models of successful regime changes, but, as Fukuyama notes, those cultures were highly developed prior to WWII.  Instead, the professor offers American ventures in Cuba, the Philippines, the Dominican Republic and Haiti as more relevant, noting that in neither instance did our intervention result in timely and effective democratization.

Intercession becomes more promising, Fukuyama argues, when the democratic initiative emanates from within the target society.  Such initiative is unlikely, of course, unless the culture’s history supports it and unless the regime is already semi-authoritarian and at least somewhat tolerant of both political organization and economic liberation.

On the one hand, the author appears to recognize the obstinacy of certain religious traditions, but, on the other, he refuses to acknowledge the degree to which Islamic culture is inherently irreconcilable with the Western values of freedom of conscience, expression and economic self-determination.

While confessing to a “large number of unknowns” concerning the “nature of the terrorist threat,” including the sources from which it draws new recruits and the parameters and geographic borders of its support, Fukuyama denies that Westerners are embroiled in what either Bernard Lewis or Samuel Huntington referred to as a clash of civilizations.  “We are not fighting Islam,” the author claims, “but a radical ideology that appeals to a distinct minority of Muslims.”

Given these admittedly numerous unknowns, and in light of the consistently surly and violent history of relations between the West and Islam, upon what facts and theory does Fukuyama base this conclusion?

“Genuine Muslim religiosity,” he contends, is and has always been a “local or national” phenomenon, and not the result of radical attempts to universalize or globalize doctrine.  Thus, for Fukuyama, religion is an insignificant part of the problem, and, because Westerners have no reason to deal with Islam as such or to impose democracy through force, all we can “hope for” is that “radical Islamists” will “eventually evolve into more responsible political parties willing to accept pluralism.”

Many have argued similarly on behalf of religious moderation or even evolution, and, of course, conscientious but intractable religionists must so argue in order to feel both rational and devout.  Nevertheless, if a religion is to survive, it must, at inevitable and critical moments, fall back upon its foundational texts.  As such, perhaps both Islam and Christianity will always be dogmatic, oppressive and universalist because, when the religious meme is threatened—either from without or from within—followers will expect their religious leaders to reveal clarity and authority, and, in turn, such leaders will feel compelled to return to the Qu’ran and the Hadith, or to Deuteronomy, Ephesians and Revelations.  Thus, others will argue, Abrahamic monotheisms will remain both dictatorial and violent and, on balance, much more harmful than beneficial.

Regardless, America at the Crossroads is an informed and thoughtful illumination of neoconservative origins, principles and internal complexity and a conscientious proposal for the future of American foreign policy in the Middle East and beyond.  Better yet, Francis Fukuyama once again effectively challenges American readers to rethink their traditional political affiliations, their democracy by proxy, and to tutor themselves regarding the leading theories of international relations, theories that, unfortunately, are seldom discussed openly among politicians or the popular media.

The Evolution of Primate Altruism.

by Kenneth W. Krause.

Kenneth W. Krause is a contributing editor and “Science Watch” columnist for the Skeptical Inquirer.  Formerly a contributing editor and books columnist for the Humanist, Kenneth contributes regularly to Skeptic as well.  He may be contacted at

AS WE SOAR INTO AN INSPIRING NEW ERA OF genomics, genetic manipulation, and, potentially, the directed evolution of our own species, naturalists urge us to remain partially grounded—to keep digging for long-buried evidence of key pre-historical developments. In so doing, however, the world’s leading anthropologists and primatologists have immersed themselves in a now- roiling debate over the origins of human morality in general and altruism in particular.

Some say that altruism—sometimes referred to as “other-regarding preferences” or “unsolicited prosociality”—is nothing more than a veneer, a cultural innovation humans alone have achieved in order to collectively restrain each individual’s natural proclivity to serve only herself, her close genetic relatives, and those who have demonstrated an adequate inclination to reciprocate to her eventual benefit. For these folks, no act can be characterized as wholly unselfish.

Others argue that altruism is more primitive than culture and, in fact, considerably more ancient than the human species itself. Other-regarding preferences, they say, are deeply innate, predating even the phylogenetic split that occurred six million years ago among the common ancestors of chimps and bonobos on the one hand and all species of hominid on the other. According to this camp’s credo, selflessness is as natural as appetite.

One line of experiments has confronted the issue directly, inquiring whether non-human primates will seize opportunities to assist others. In 2005, for example, UCLA anthropologist Joan Silk and others chose 18 chimpanzees (Pan troglodytes) as the subjects of two such experiments, conducted in Louisiana and Texas.(1)  Chimps are among the primates most likely to exhibit unsolicited prosocial behavior, they reasoned, because in the wild they regularly hunt, patrol, and mate-guard cooperatively.

In each study, subject chimps were allowed to deliver food to other chimps, or “conspecifics,” at no cost to themselves. The test apparatuses provided each confined subject with two options—the 1/0 choice where it could acquire food only for itself, and the 1/1 choice where it could obtain food for both itself and its separately caged partner. As an essential control, acting chimps were given the same options with no partners present.

Silk’s team predicted that if chimps are truly altruistic they should choose the 1/1 option more often than the 1/0 option when a conspecific is there to benefit. But that wasn’t the case. In Louisiana, not one of the seven subjects chose the 1/1 option significantly more often when partnered. In Texas, the remaining 11 actors went with both the 1/1 and the 1/0 option an average of only 48% of the time when another chimp was present.

“The absence of other-regarding preferences in chimpanzees,” the authors inferred, “may indicate that such preferences are a derived property of the human species tied to sophisticated capacities for cultural learning, theory of mind, perspective taking and moral judgment.” Nevertheless, Silk’s team remained open to the prospect that altruism might be detected among primates that, in some crucial ways, were even more cooperative than chimps. We will consider that possibility later.

Altruism’s Alter-Ego.

A closely related line of experiments has tackled the same issue from a different direction, asking instead whether primates display a rudimentary sense of fairness in some form of “inequity aversion” (IA). If an animal reacts negatively to its own relative overcompensation, we say it has demonstrated some sensitivity to “advantageous inequity.” If it merely responds to a conspecific’s superior gain, on the other hand, the animal has shown aversion only to “disadvantageous inequity.”

The former inclination probably evolved after (and, morally speaking, is emphatically more advanced than) the latter because an animal sensitive to its own advantage can demonstrate not only an egocentric expectation of how it should be treated, but also a communal expectation of how all members of its species should be treated. In either case, if test subjects attempt to restore equity by sacrificing their own gains—even if only to simultaneously and unceremoniously deny superior gains to their luckier partners—according to many (but not all) researchers, they have nonetheless acted altruistically.

In 2003, Emory University primatologists Sarah Brosnan and Frans de Waal developed token exchange experiments where tufted capuchin monkeys (Cebus apella) were measured for their reactions to situations in which their partners received greater food rewards.(2) In the end, shortchanged subjects proved less likely to complete exchanges for identical tokens, and withdrew even more frequently when their partners received prizes for no tokens at all. These now-classic results have been widely interpreted as formidable evidence of disadvantageous IA in primates.

Two years later, Brosnan, de Waal, and Hillary Schiff released the outcomes of a similar study of adult chimpanzees.(3) In order to distinguish the effects of social alignment, the team chose four animals that had lived continuously in pairs and 16 others that had been housed together at the Yerkes National Primate Research Center in Atlanta, Georgia for either 30 years or eight years prior to testing. As in the 2003 experiment, subjects were given tokens—in this case, rather useless and nondescript chunks of white PVC pipe—which they had been trained to return for either cucumber slices (the low-value reward) or grapes (the high-value reward).

During the inequity test, examiners initially allowed the partner chimps to exchange for a juicy, delicious grape—while eager subjects observed, of course—and then offered the subjects a relatively dry and no doubt disappointing cucumber slice. The examiners diligently recorded the subjects’ reactions, noting whether they had accepted or rejected their prizes. Brosnan discovered first that, when the tables were turned, subjects did not react negatively when given a superior reward and, thus, were likely not averse to advantageous inequity. Whether such a finding actually distinguished chimpanzees from humans in any meaningful way, the authors noted, was questionable.

Second, according to Brosnan, the results confirmed that disadvantageous IA was “present and robust” among chimpanzees, although to significantly different degrees depending on each subject’s social history. Chimps that had lived in pairs or in relatively novel groups reacted most intensely, while animals from older, more tightly-integrated groups appeared more accepting of inequity—all of which could be entirely consistent with human predilections to either “make waves” or “go with the flow,” depending primarily on their social milieu. Tolerance of inequity, Brosnan suggested, may be more a function of group size and intimacy than either moral choice or any isolated cognitive factor.

So by the end of 2005, very little if anything had been truly settled. The experiments would continue and become ever more creative and exacting, but the already muddied anthropological waters would grow more cluttered and murkier still.

High Expectations.

In 2006, three teams from the Max Planck Institute for Evolutionary Anthropology in Leipzig, Germany published studies that in one way or another challenged these landmark outcomes from Yerkes. Julian Brauer’s group tested for IA among chimps, bonobos, gorillas, and orangutans—30 individual great apes in all—and produced a pattern of food rejection that was opposite to that reported by Brosnan.(4) In other words, instead of snubbing more food after seeing their partners receive tastier treats, Brauer’s apes actually rejected less food.

All the same, the authors did not infer that apes were necessarily insensitive to unfairness. In fact, at one point they questioned whether food refusal was a fair test of IA to begin with. Inequity-wary apes, after all, might decide to accept lower-quality spoils simply in an effort to offset the higher quality gifts bestowed upon their partners. Citing a then-recently published study questioning chimpanzee altruism, however, Brauer’s team finally betrayed a clear inclination to attribute their results to the so-called “food expectation hypothesis,” which asserts that the mere act of witnessing a conspecific’s receipt of superior food will create an anticipation of acquiring the same food for oneself. Such an expectation might explain why Brauer’s chimps begged more vigorously and why many of her apes generally remained at their testing stations much longer after having witnessed partner overcompensation.

The study Brauer cited had been conducted by a second German team led by Keith Jensen.(5) The key problem with Brosnan’s examination, according to Jensen, was that subject chimps were never allowed to convey their mindsets by actually correcting unequal outcomes. Silk’s group had devised a somewhat more effective experiment in this respect, the authors commended, but even they had failed to test for any- thing more than selfishness (the 1/0 option) or mutualism (the 1/1 option). In Jensen’s experi- ments, by comparison, 11 chimps participated in three separate studies collectively designed to reveal expressed tendencies toward altruism and spite as well.

In study one, each subject was allowed to pull one of two tables toward itself. The first table contained bananas accessible to both the subject and its partner; the second table held fruit accessible only to the subject. Either way, subjects received the same reward. But because Jensen’s chimps predominantly chose the mutually accessible table in both the test (partner present) and control (partner absent) conditions, the results were inconclusive as to selfishness and mutualism. Nevertheless, Jensen asserted, this initial phase of the experiment did show that his chimps were not averse to disadvantageous inequity, at least with regard to relative effort expended.

Studies two and three tested for spite and altruism. In neither case could any subject receive a reward for pulling any table closer to itself or its partner. In experiment two, acting chimps could have conveyed an other-regarding preference by pulling tables accessible only to partners, or passive spite by doing nothing at all more frequently in the test condition than in the control condition. But they did neither.

In experiment three, the scheme was altered slightly such that, in order to deny food to their partners, subjects needed to actively draw those partners’ trays away. But, once again, the Leipzig chimps were as likely to do nothing in one condition as in the other, thus failing to demonstrate active spite as well. The authors noted, however, that two of their six chimps did express potential signs of altruism. But these animals also tended to beg or harass their partners following delivery of the fruit, thus raising the possibility that they intended to only benefit themselves.

Food for Thought.

Felix Warneken and Michael Tomasello, also from the Planck Institute, decided to examine the issue through an entirely different type of experiment. If altruism exists among our primate cousins, they judged, it might more readily be elicited with something less critical to individual survival than food. So Warneken and Tomasello tested both human children (24 18-month-old infants) and three young chimpanzees (34, 54, and 54 months old) for their willingness to help human caretakers (quite familiar to the chimps) with some task absent of any possible expectation of reward.(6)

As predicted, the children assisted experimenters more often and in a greater variety of tasks than the chimps. Nonetheless, each of Warneken’s chimps reliably helped a reaching human obtain apparently desired objects. Although young humans clearly cooperate to a degree found in no other species, the authors concluded, “our nearest primate relatives show some skills and motivations in this direction as well.”

By the summer of 2007, Warneken had assembled another team and published the results of similar “instrumental helping” experiments calculated to address several important and yet unanswered questions—in particular, whether 36 semi free-ranging chimps would spontaneously help unfamiliar humans and genetically unrelated conspecifics in addition to their caretakers, and whether they would do so at some significant cost to themselves.(7)

Again, as expected, infant children helped more quickly. But the chimps performed just as reliably regardless of their partners’ familiarity or species, even when they had to expend a little extra effort to do so. “The roots of human altruism may go deeper than previously thought,” Warneken ultimately concluded, “reaching as far back as the last common ancestor of humans and chimpanzees.”

Later that year, however, Keith Jensen’s team cast a skeptical eye on Warneken’s conclusions in two well-focused examinations of potential IA among chimps. The first pair of experiments probed 11 animals’ capacities for spite—altruism’s evil twin, if you will.(8)  Jensen reasoned that chimps might be characterized as altruistic, at least in a punitive sense, if they choose to act out against conspecifics due to an abstract sense of fairness. In the first study, caged subjects were allowed to pull ropes to collapse food-laden tables drawn away by humans toward different enclosures that either contained other chimps (test condition) or were empty (control condition). In the end, the chimps appeared to be indifferent. Actors were just as likely to collapse the platforms when they approached empty cages as when they neared hungry conspecifics.

In the second study, subjects were exposed to three conditions. In conditions one and two, much as before, human experimenters pulled the tables away from subject animals and toward partners or empty cages. In the last condition, however, it was the partner chimps that were allowed to drag the tables away from subjects. Between the first two conditions there was no real disparity, indicating again that subjects didn’t really care whether their partners benefited inequitably. Between the first two and the third conditions, on the other hand, subjects were significantly more likely to drop the tables when other chimps, as opposed to humans, began drawing them away. From these combined results, Jensen concluded that although chimps are certainly vengeful, “[s]pitefulness may thus be a peculiarly human phenomenon.”

Ultimatums (and more Expectations).

Hailing it as the “benchmark test for examining sensitivity to fairness and other-regarding preferences,” Jensen then unleashed his 11 subjects on a chimp-friendly version of the celebrated ultimatum game.(9) Proposer animals were permitted to make one of two possible offers to their receiving partners, potentially retaining either 100, 80, 50, or 20% of the spoils for themselves in each trial. If the receiver accepted the offer, each party got what it wanted. But if the receiver rejected the offer—having noted what the proposer intended to keep for itself—neither animal received any reward.

Presumably out of some concern for fairness, humans proposers tend to make equitable offers of 40 to 50 percent or, as receivers, to reject offers of 20 percent or less, thus confounding the economic model of rational self-interest (so-called Homo economicus). This was not how the Leipzig chimps reacted, however. Proposers chose not to make fair offers and receivers opted to accept all nonzero offers without hesitation or perceptible sign of irritation. While the authors cautioned that these outcomes “may be in part be a reflection of the fact that active food sharing is rare among the species,” they were clearly inclined to attribute such behaviors to the chimps’ absent sense of justice.

In late 2007, Megan van Wolkenten, working with Brosnan and de Waal, finally published a narrowly tailored response to Brauer and others addressing the alleged preeminence of food expectation over IA.(10) They used the now-familiar token exchange experiment—this time enhanced with an additional condition where food rewards were shown to subjects well prior to exchange— on 13 capuchin monkeys. But, contrary to the predictions of various expectation hypotheses, behavioral changes did not depend on either greed or frustration. Rates of refusal among subjects, in fact, increased not when higher value grapes were merely visible, but only when they were actually bestowed upon partners.

Importantly, van Wolkenten’s subjects also made significantly fewer exchanges when forced to expend more effort for the same lower value cucumbers received by partners. As the food value increased, however, effort became secondary, indicating that capuchins are willing to reprove inequity only when the cost of doing so is slight. This appreciable yet limited brand of IA, the authors proposed, “likely evolved in conjunction with cooperative enterprises,” and “may characterize a great variety of social animals.”

More Monkey Business.

By the end of 2007, then, the combined body of research had established mixed results at best, especially with regard to the great apes. Recalling Joan Silk’s suggestion that true altruism might be discovered among primates even more social than chimpanzees, Swiss anthropologist Judith Burkhart’s team decided to test 26 common marmoset monkeys (Callithrix jacchus) in two studies—one for related, one for unrelated pairs —involving hungry partners and subject operated food trays.(11)

The experimenters provided each actor with a 0/0 option and a 0/1 option only, thus elimi- nating all potential for subject rewards. Because marmosets are cooperative breeding callitrichids, Burkhart predicted that if any primate should display an unsolicited prosocial tendency capable of overcoming any penchant for envy, it would be this species, despite their theory of mind deficit and general cognitive shortcomings.

Burkhart was right. Kin or no kin, marmoset subjects—fully schooled with the test apparatus and, thus, aware of the experiments’ conse- quences—pulled the 0/1 tray more often when their partners were present in adjacent cages than in the control condition when their partners were absent. Remarkably, the disparity widened significantly when female “helpers”—which, despite this label, tend not to carry other monkeys’ infants in the wild—were eliminated from the analysis. Because humans and New World monkeys are the only primates that behave as cooperative caretakers, Burkhart proposed, strong altruism may have evolved within such groups independently, and not necessarily among the ancestors common to chimps, bonobos, and humans.

More Food for Thought.

The thick, swirling waters of controversy have spilled largely unabated into 2008. Working with Brosnan, Silk, and others, American evolutionary psychologist Jennifer Vonk published a detailed study of low-cost, conspecific-directed altruism among 18 chimpanzees at the University of Louisiana’s Cognitive Evolution Group laboratory.(12) In two separate experiments involving two different apparatuses and two distinct groups of chimps, actors were given the options to trigger rewards for themselves alone, for their partners alone, or for both themselves and their partners.

The team chose these three options in order to address important criticisms of previous experiments involving food. Because her chimps were allowed to act prosocially only after having fed themselves, Vonk argued, this method avoided the possibility that subject animals might be distracted from an otherwise spirited altruistic tendency by the potent and ever-present need to feed.

If chimps are really other-regarding, the authors reasoned, subjects should deliver rewards to partner enclosures at some point during the experiment, but more often in the partner-present test condition than in the partner-absent control condition. By contrast, if chimps are indifferent to the welfare of others, actors should minimize their personal costs by obtaining rewards only for themselves. Ultimately, the presence of awaiting partners in other enclosures had no significant effect on subjects in either experiment. At first, actors consistently released both rewards. But delivery rates to other cages always decreased as subjects learned that such efforts would not benefit them.

Notably, one of the 11 chimps tested in the second experiment did choose to act prosocially, but these results could not be replicated. “[W]hile chimpanzees’ behavior is consistent with standard evolutionary models based on kinship and reciprocity,” Vonk insisted, “human cooperation and prosociality may require an emerging class of evolutionary models, rooted in the coevolutionary interaction of genes and culture.”

Agreeing to Disagree.

Despite these equivocal results, some scientists still see altruism as a considerably more ancient impulse, born of the intense parental and, thus, empathic instinct. Frans de Waal, as one promi- nent example, appears to be thoroughly convinced that some skeptics of primate altruism have their arguments backwards—at least in one crucial respect. “[E]mpathy evolved in animals as the main proximate mechanism for [individually] directed altruism,” he explained in a recent review, and it is empathy—not self-interest—that “causes altruism to be dispensed in accordance with predictions from kin selection and reciprocal altruism theory.”(13) Although gene propagation and benefit exchange may be the evolutionary or ultimate cause of altruism, only a spontaneous emotional response to another being’s situation can possibly trigger or proximately cause an altruistic impetus.

In his latest study of non-cooperatively breeding monkeys, de Waal discovered that brown capuchins will predominantly choose the 1/1 mutual option over the 1/0 selfish option, depending on the subjects’ familiarity with their partners.(14) Although his monkeys’ other-regarding tendencies clearly turned on social closeness, de Waal nevertheless concluded that because kinship was critical and because his subjects had no means of predicting return favors, only empathy could explain this study’s results.

When I asked him about the persisting debate, de Waal proposed that the scientific community has become polarized between evolutionary biologists on the one side and, on the other, a discrete group of economists and anthropologists that “has invested heavily in the idea of strong reciprocity,” which absolutely demands discontinuity between humans and all other animals. As for the results obtained by Silk and others, de Waal offered, experiments such as these involving repeated trials and frequent rewards are vulnerable to “side-biases” that can skew outcomes.

Sarah Brosnan, a former student of de Waal’s and now Assistant Professor of Psychology at Georgia State University, remains ambivalent. Her subjective though surely copious experience with both apes and monkeys informs her that at least some of these animals do seem altruistic. Even so, she told me, “there is not too much evidence for this outside some of Frans’ and Felix Warneken’s work.” But cooperation in all species, she emphasized, “is much more likely to be based on emotion and relationships than on cognitive calculations.”

Both de Waal and colleague Keith Jensen are doubtful that even chimpanzees possess the cognitive capacities requisite for delayed reciprocation. But for Jensen, the added conclusion that chimps must be altruists simply doesn’t follow. “De Waal’s use of the term ‘empathy’ is somewhat contentious,” he told me, “and the evidence he provides for empathy is [anecdotal and] not very robust.”(15) More evidence is needed, he admitted, but, like Jennifer Vonk, his “working hypothesis” is that other-regarding preferences emerged at some point during human evolution only.

Even so, both Brosnan and Jensen conceded that the distinction between food exchange and instrumental helping is a potentially crucial one. Indeed, Jensen and Felix Warneken are now collaborating on a new project to determine whether food rewards might interfere with genuine other-regarding preferences. Although “food exchange is not a bad test for altruism,” Warneken reminded me, it explores “only one type of potentially altruistic behavior.” In the more sensitive context of instrumental tasks, he added, chimpanzees have repeatedly demonstrated solid helping tendencies.

When I asked Warneken about Vonk’s latest attempt to neutralize the nutritional imperative, he warned that Vonk’s chimps might not have fully understood how the apparatuses worked during that experiment’s altruism phase. “The pattern of results,” he argued, “still suggests that the subjects had a tendency to try to obtain the reward for themselves.” Plugging Jensen’s 2006 study as the most convincing presentation to date of limited prosociality among chimps, Warneken recommended that future researchers follow that team’s lead, at least with respect to designing an apparatus that animals might comprehend more intuitively.

Where to Go from Here.

Everyone agrees that more work needs to be done, and that no research could be more germane to achieving a competent grasp of who we are as a species and where we might be headed. If altruism is in fact deeply innate to humanity’s collective being, we may have to rethink a number of things, including some of our most established political and economic assumptions.(16) Jensen summed it up pretty well when I invited him to characterize his work’s significance:

This research is interesting to the question of what makes humans special, if, indeed, they are. Most research in the past has focused on “cold cognition” such as abstract reasoning, language and tool use. Social motivations and emotions—“hot cognition”—are just as important, and may even be central to the emergence of human ultrasociality. Holding a lens up to ourselves after focusing it on other species will help us see ourselves more clearly.

So it looks like we’ll be hearing a great deal more from these and other esteemed researchers during the coming years. Sadly, however, the indispensable subjects of these investigations seem to be living on borrowed time, the African great apes especially. If scientists can ever clear the dim, shadowy depths of altruistic origins, they’ll have to act quickly before our own dark natures drive our ancestral cousins into extinction.


  1. Silk, J. B., Brosnan, S. F., Vonk, J., Henrich, J., Povinelli, D. J., Richardson, A. S., Lambeth, S. P., Mascaro, & Shapiro, S. J. 2005. “Chimpan- zees Are Indifferent to the Welfare of Unrelated Group Members.“ Nature, 437, 1357-1359.
  2. Brosnan, S. F. & de Waal, F. B. M. 2003. “Monkeys Reject Unequal Pay.” Nature, 425, 297-299.
  3. Brosnan, S. F., Schiff, H. C. & de Waal, F. B. M. 2005. “Tolerance for Inequity May Increase With Social Closeness In Chimpanzees.” Proc. R. Soc. B, 272, 253-258.
  4. Brauer, J., Call, J. & Tomasello, M. 2006. “Are Apes Really Inequity Averse?” Proc. R. Soc. B, 273, 3123-3128.
  5. Jensen, K., Hare, B., Call, J. & Tomasello, M. 2006. “What’s in it for me? Self-regard Precludes Altruism and Spite In Chimpanzees.” Proc. R. Soc. B, 273, 1013-1021.
  6. Warneken, F. & Tomasello, M. 2006. “Altruistic Helping In Human Infants and Young Chimpanzees.” Science, 311, 1301-1303.
  7. Warneken, F., Hare, B., Melis, A. P., Hanus, D. & Tomasello, M. 2007. “Spontaneous Altruism By Chimpanzees and Young Children.” PloS Biology, 5(7), e184.
  8. Jensen, K., Call, J. & Tomasello, M. 2007. “Chimpanzees Are Vengeful But Not Spiteful.” Proc. Natl. Acad. Sci., USA, 104, 13046-13050.
  9. Jensen, K., Call, J. & Tomasello, M. 2007. “Chimpanzees are Rational Maximizers In an Ultimatum Game.” Science, 318, 107-109.
  10. van Wolkenten, M., Brosnan, S. F. & de Waal, F. B. M. 2007. “Inequity Responses of Monkeys Modified by Effort.” Proc. Natl. Acad. Sci., USA, 104, 18854-18859.
  11. Burkhart, J. M., Fehr, E., Efferson, C. & van Schaik, C. P. 2007. “Other-Regarding Preferences In a Non- Human Primate: Common Marmosets Provision Food Altruistically.” Proc. Natl. Acad. Sci., 104, 19762-19766.
  12. Vonk, J., Brosnan, S. F., Silk, J. B., Henrich, J., Richardson, A., Lambeth, S.,Schapiro,S. & Povinelli,D. J. 2008. “Chimpanzees Do Not Take Advantage of Ver y Low Cost Opportunities to Deliver Food to Unrelated Group Members.” Animal Behavior, 75, 1757-1770.
  13. de Waal, F. B. M. 2008. “Putting the Altruism Back Into Altruism: The Evolution of Empathy.” Annu. Rev. Psychol., 59, 279-300.
  14. de Waal, F. B. M., Leimgruber, K. & Greenberg, A. R. 2008. “Giving Is Self-rewarding for Monkeys.” Proc. Natl. Acad. Sci., USA. 105, 13685- 13689.
  15. See also, Silk, J. B. 2007. “Empathy, Sympathy, and Prosocial Preferences In Primates.’ In: The Oxford Handbook of Evolutionary Psychology. (Ed. by R. I. M. Dunbar & L. Barrett), pp. 115-126. Oxford: Oxford University Press (“Current claims for the existence of empathy, sympathy, moral sentiments, and other-regarding preferences in other primates rest on an insecure empirical foundation.”).
  16. See, e.g., Bowles, S. 2008. “Policies Designed for Self-interested Citizens May Undermine ‘the Moral Sentiments’: Evidence from Economic Experiments.” Science, 320, 1605-1609 (“Economists, psychologists, and others … are well on their way to constructing an economic psychology of the interplay of self-regarding and other-regarding motivation that may eventually enlighten mechanism design and public policy.”).