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With their 1991 publica-tion in The QuarterlyReview of Biology, TheDawn of Darwinian

Medicine,1 George C. Williams, anevolutionary biologist, andRandolph Nesse, an evolutionarypsychiatrist, essentially establishedthe foundation for a new subject tobe incorporated into the medicalschool curriculum; EvolutionaryMedicine (EM), also referred to asDarwinian Medicine, are terms usedto describe a new paradigm inmedical education that attempts tounderstand modern diseasesthrough application of evolutionarytheory and human ecology.

Over the past 20 years, thesubject of Evolutionary Medicinehas been gradually emerging acrossNorth America and is now quickly

growing throughout Europe andother parts of the world into a legit-imate academic discipline.Presently, there are several text-books, peer-reviewed scientific arti-cles, websites and blogs, majorinternational symposiums, medicalschool curriculum modules andadvanced post-graduate courses ofstudy, all dedicated to this excitingnew field of scientific inquiry(Appendix-The Evolutionary &Medicine Review). It appears EM ishere to stay for, the medical profes-sion….but what about the rest ofthe allied-health professions?

Evolutionary DentistryIn the spring of 2012, the

National Evolutionary SynthesisCenter (NESCent) will host a ‘cataly-sis meeting’ that will bring together

clinicians and researchersfrom several disparate fields(e.g., evolutionary biology,paleopathology, biomechan-ics and food science) toexplore the implications ofthe evolution of humanteeth and jaws for dentistryand orthodontics. Theprogram, “EVOLUTION OFHUMAN TEETH AND JAWS:Implications forDentistry andOrthodontics”, willexplore the ideathat many of ourcurrent dentaland orthodonticproblems relateto a mismatchbetween thechemical andphysical proper-ties of the foods

we eat today, and those to which ourancestors’ jaws and teeth had beenbest adapted.

The subject of Evolutionary OralMedicine (EOM) as a proposedacademic discipline within the fieldof dentistry, was recently introducedat The Ancestral Health Society’s(AHS) First Symposium on AncestralHealth held at UCLA in August 2011by Kevin Boyd, a pediatricdentist/nutritionist who is currentlystudying Biological Anthropology,and Michael Mew, an orthodontistin the UK who is interested in EOMas it pertains to understanding theetiology of malocclusion (Appendix-Y Crooked Teeth).

Consistent with the aforemen-tioned NESCent program theme, theirpresentation, “Where is Darwin onDentistry? Caries and Malocclusionfrom an Evolutionary Perspective,2centered around the observation thatdental caries and malocclusion, whilenow highly prevalent public healthdiseases, are both surprisingly rarewithin the pre-Industrial skeletal andpre-historic fossil records, and alsoseldom seen in many present-day non-westernized cultures.3, 4, 5 (Figs.1, 2 and3) According to Profit,6 the fact thatmalocclusion now occurs in a major-

By Kevin L. Boyd, M.Sc., DDS

DARWINIANDENTISTRYPART 1:

DARWINIANDENTISTRYPART 1:

Fig. 1

Fig. 2

An Evolutionary Perspective on the Etiology of Malocclusion

Indian skulls studied by Dr. Weston A. Price. Each skull hasnice occlusion and no decay. Adapted from Palmer, 2003.(Copyright© Price-Pottenger Nutrition Foundation®)

70,000 year old skull with nice occlusion and no decay.Adapted from Palmer, 2003.

34 November/December 2011 JAOS

ity of the population does not meanthat it is normal; skeletal remainsindicate that the present prevalencesare several times greater than it wasonly a few hundred years ago. Crowd-ing and malalignment of teeth wasunusual until relatively recently, butnot unknown.

Furthermore, other than somefossil evidence of anterior incisorcrowding in ancient Egyptian skulls,it is unlikely that class II and classIII skeletal malocclusion appearedappreciably in humans until aroundthe time of the Industrial Revolu-tion in Western Europe during themid-18th century,7 and whendetected, it was usually confined toprivileged-class individuals. Dentalcaries on the other hand has beenplaguing mankind since the adventof agriculture some 10,000 yearsago, and there is even fossilevidence of tooth decay as far backas 1.5 million years ago in one (pre-human) Paranthropus robustus skull.8

A sharp rise in caries prevalence,however, doesn't appear in modernhumans until nearly 1,000 years ago(Fig.4) with the introduction of canesugar to Western Europe,8 and onlybegan to reach epidemic proportionsin the late 19th/early 20th centuries.Susceptibility to dental caries, clearly adietary-infectious disease caused byincreased sugar consumption resultingin increased acid production by oralbacteria, is not likely influenced by

large genetic changes that might haveoccurred since the Agricultural Revolu-tion some 10-15,000 years ago.

While the cause of malocclusionis less clear, it is also not likely aresult of recent genomic change.This article will develop the hypoth-esis that malocclusion, when viewedfrom an EM perspective, resultsfrom a consequential mismatchbetween a stone-age adapted masti-catory apparatus, and a post-Indus-trial feeding environment.

Evolutionary Biology;Basic Science

According to Nesse9 “…fewphysicians and medical researchershave taken a course on evolution-ary biology, and no medical schoolteaches evolutionary biology as abasic science for medicine. It is as ifengineering students never learned

physics.” Perlman10 suggests thatthe reason for this void in pre-medical and medical school educa-tional requirements is related to theorigins of the framework for thecurrent medical didactic model, theFlexner Report.11

At the behest of the AmericanMedical Association in the early1900s, the highly respected Americaneducation researcher, AbrahamFlexner, was given a commission bythe Carnegie Foundation to reformmedical education; at that time, evenwithout a high school diploma onecould essentially buy a medical degreeafter serving an apprenticeship inmuch the same manner as any othertrade school of that era. The AMA, a

www.orthodontics.com November/December 2011 35

Fig. 3

Fig. 4

Number of carious teeth per 100 teeth in fourEuropean populations. Adapted from Kean, 1980.

“According to Profit,6

the fact thatmalocclusion now

occurs in a majority ofthe population does not

mean that it is normal;skeletal remains

indicate that thepresent prevalences are

several times greaterthan it was only a few

hundred years ago.”

Prehistoric skull with normal palate, wide dental arch, and large posterior nasal aperture. Note the U-shaped arch (left). Younger, “modern” skull(1940s) with a high palate, narrow dental arch, and small, congested posterior nasal aperture. Adapted from Palmer, 2003.

36 November/December 2011 JAOS

relatively weak organization then,recruited Flexner to help elevatemedical educational institutions tothe standards of a few US schools, likeJohn’s Hopkins, Michigan andHarvard that followed the Germanmodel for physician training. Accord-ing to Perlman, “Evolutionary biologywas a poorly developed discipline atthe time of the Flexner Report andwas not included in his recommenda-tions for premedical or medicaleducation….”. Diller12 recentlystated, “America and the medicalprofession desperately need a newFlexner Report for the 21st century.”

Following the AMA and Flexner’slead, in mid-1920s, William Gies, aColumbia University biochemistryprofessor and future founder ofthe Journal of Dental Research,at the behest of a consortiumof university-affiliated dentalschools, also received acommission from Carnegie tohelp elevate dental education;dental schools in the 1920ssuffered from many of thesame problems as pre-FlexnerReport medical schools—theywere mostly low-grade tradeschools unaffiliated withuniversities. In 1926, the GiesReport13 on Dental Educationin the United States andCanada was released as afollow-up to the 1910 Flexner,and similarly, did not includea recommendation for includ-ing courses in evolutionary biologyinto the didactic curriculum.

In reference to the present didacticcurriculum in dental education,Baum14 states, “… many changes havebeen made since then, but the basicdesign and approach remain thesame.” Dr. Baum also argues that,while dentistry has benefitted tremen-dously from the findings containedwithin the 1926 Geis Report, “…weshould be mindful that it was writtena full 80 years ago. At that time, thebiological sciences were much moreprimitive and phenomenological, thepopulation had very different kinds ofdental problems…”.

Evolution andVulnerability to Disease

In their 1994 book, Why We GetSick: The New Science of DarwinianMedicine, Nesse and Williams15

describe how bodies have traits thatcan often leave them vulnerable todisease for a variety of reasons: ‘co-evolution with pathogens’ and‘genomic mismatch with themodern environments’ are at thetop of their list; other explanationsinclude ‘trade-offs’, ‘constraints onnatural selection’, ‘reproductivesuccess at the expense of health’and ‘protective defenses that areeasily confused with diseases’.

Dental CariesThe cariogenic group of bacteria

most commonly implicated in dentalcaries, Mutans streptococcus (MS) hasbeen co-evolving with humans sincewe began migrating out of Africa tens

of thousands of years ago. In linewith Nesse and Williams’ ‘human co-evolution with pathogens’ hypothe-sis, Caufield16 suggests that, similar tomitochondrial DNA, genetic mappingof MS’ DNA could represent a ‘secondgenome’ that might someday be usedto verify early human migratorypatterns throughout the world. Addi-tionally, the ‘genomic mismatch withthe modern environments’ hypothe-sis might also be a good explanationfor why dental caries is only a rela-tively recent finding in humanhistory that also seems to coincidewith the first appearance of refinedgrains and sugars in the diet.

Malocclusion

Anthropologists have longreported that human craniofacialvolume has been diminishing sincethe advent of agriculture in the

Middle East, and most rapidly sincethe early/mid-18th century in West-ern Europe: Larsen17 reports, “…ashift to agriculture or more intensi-fied agriculture was accompanied byan increase in dental crowding andmalocclusion.”; Gilbert5 states, “…jaw anomalies (malocclusionswherein the teeth cannot fit prop-erly in the jaw) are relatively new toEuropean populations.

Well-preserved skeletons from the15th and 16th centuries showalmost no malocclusion in thepopulation….”; and Lieberman18

reports in his recent book, Evolutionof the Human Head, “….there ismuch circumstantial evidence thatjaws and faces do not grow to the

same size that they usedto…”.

Corruccini19 has hypothe-sized, “An epidemiologictransition to high prevalenceof such diseases as diabetesand coronary heart diseaseaccompanies the process ofmodernization/industrializa-tion. I suggest that anequally clearly definedepidemiologic transitioncharacterizes malaligned anddiscrepant dental occlusal

relations in western soci-eties, and others undergo-ing urbanization, and thatthe rapidity of the transi-tion is proportional to therapidity of urbanizational

change. This phenomenon ratherthrows the weight of suspiciontoward environmental, not genetic,etiologic factors.”

Anatomically ModernHumans & theMasticatory Apparatus

Fossil evidence from East Africais consistent with recent geneticevidence indicating that modern-day humans (Homo sapiens) havelikely been in their presentanatomic form (AnatomicallyModern Human/ AMH-Fig.5) forapproximately150,000-200,000years;20 the first evidence of modernEuropean (Fig. 6) Homo sapiens(Cro-Magnons) can be dated toapproximately 35,000 years ago.21

This implies that the genomecoding for the modern anatomicform (phenotype) of our ancient

Fig. 5

Fig. 6

Artist’s reconstruction of a 160,000 year-old AMH. Adaptedfrom Sanders, 2003.

Artist’s reconstruction of a 35,000year-old Cro-Magnon. Adapted fromAmerican Museum of NaturalHistory, 2011.

www.orthodontics.com November/December 2011 37

African ancestors, is also very little,if at all, changed over thousands ofgenerations. This has significantimplications for why we arepresently experiencing such highprevalence of malocclusion over thepast few hundred years… Macroge-nomic change has not been shownto occur over such a relatively shorttime span.

The component of the craniofa-cial complex that is dedicated tothe function of initiating mechani-cal/ chemical processing of foodprior to its subsequent digestionand assimilation of nutrients forlater biological utilization, is calledthe human masticatory apparatus(HMA). As our pre-modern humanancestors evolved away from theircommon ancestor with the modernchimpanzee over 6 million yearsago, the HMA had to have beenindispensible to their ultimateevolutionary success. Thus, thecombination of genes involved incoding for the modern HMA-phenotype is likely little changedsince modern Europeans firstappeared nearly 35,000 years ago,and many anthropologists specu-late that our complete genome hasprobably not undergone anyMacroevolutionary change since wefirst appeared in Africa as AMH’snearly 200,000 years ago.

Mismatch to Modernity& Malocclusion

As Corruccini implies, the rela-tively recent appearance of maloc-clusion in humans is not bestexplained as resulting from a recentand anomalous Macroevolutionarygenomic change...that would requirea vast amount of geological time. Amore plausible explanation wouldbe one that is consistent with Nesseand Williams’ ‘Mismatch’ explana-tion for disease vulnerability; anunchanged ancient genome exposedto a less-challenging modern feedingenvironment (since the IndustrialRevolution) is now the leadinghypothesis for understandingmalocclusion etiology that isaccepted by many anthropologistsand anthropologically-informedorthodontic clinicians andresearchers…but seemingly at oddswith current orthodontic teachingand clinical practice.

In the most recently publishededition of the widely-used textbookContemporary Orthodontics,6 coau-thor William Proffit posits the ques-tion, “Is it possible that a child’smasticatory effort plays a majorrole in determining dental archdimensions?”; and then providesan answer, “That seems unlikely.”Dr. Proffit’s conclusion might be atodds with the observation thatancestral-type infant/early child-hood feeding environments (breast-feeding at-will into the third yearof life and weaning to fibrous/firm-textured first foods) seems toprovide some protective benefitagainst the later development ofmalocclusion in pre-Industrialized,prehistoric and non-Westernizedmodern-day cultures. Furthermore,there are multiple studies22, 23, 24

that clearly indicate a negativeeffect of bottle-feeding versusbreastfeeding with respect to laterdevelopment of anterior open-bitesand/or posterior crossbites.

A Need for ChangeAnthropological Norm (AN) is a

concept that is currently beingexplored by several disciplines inhealthcare;25 for example, it is nowroutine to look at free rangingSouth African Bushman, Khoisanpeople, and others for an "anthro-

pologically normal"/ prehistori-cally "natural" level of serumcholesterol, LDL to HDL ratio,blood pressure, sodium, bloodsugar. All these variables are atunnaturally high levels in modern-ized/westernized populations.

Hypothetically, AN implies theexistence of a pre-Industrial pheno-typic range for a variety of physi-cal/physiological phenotypic traits(e.g., the human masticatory appara-tus, salivary pH, etc.) that are normalfor assuring maximum survival,thriving and reproductive fitness.The AN hypothesis is predicated onthe observation that the humangenome is best adapted to pre-Indus-trial diets, lifestyles and environ-ments as it (the human genome) hasundergone virtually no Macroevolu-tionary change in perhaps the last60,000-200,000 years. As the allelesthat code for the human masticatoryapparatus are likely unchanged forthousands of generations, to suggestrevision of current “anthropologi-cally” uninformed cephalometricnorms, which are almost entirelybased upon 20th-century skulls, doesnot seem unreasonable.

In 1981, a paper by James McNa-mara appeared in the Angle Orthodon-tist26 describing a study showing thatmost of the skeletal Class II maloc-clusion subjects in a cohort of 8- to

Fig. 7

Notice the forward position of A-point relative to the N-perpendicular to the Frankforthorizontal plane; this is a common finding in pre-Industrial skulls but would indicate an abnormally protrusive maxilla by most currently used cephalometric analyses. Adapted fromCorruccini 1989.

10-year-olds were not maxillaryprotrusive, rather, most had retrusivemaxillas; the conclusions regardingrelative maxillary skeletal retrusion inthe A/P dimension were based upontwo pre-treatment cephalometricangular values: 1.) SNA (Steiner)angles-less than 81 degrees; and 2.)the distance of A-point from Nasionperpendicular (less than 0mm).

When utilizing McNamara’s A-point to N-perpendicular cephalo-metric angular measurement, pre-Industrial, prehistoric and pre-West-ernized skulls are somewhat maxil-lary protrusive (Fig. 8), these dataseem to be at least circumstantiallysupportive of the hypothesis thathuman malocclusion is a relativelyrecent phenomenon since techno-

logical advances stemming from theIndustrial Revolution in WesternEurope.Furthermore, because theywere largely developed from early20th-century (post-Industrial)databases, currently used orthodon-tic cephalometric normative valuesshould now be revised as they likelydo not represent anthropologically-accurate ideals for true genomiccraniofacial growth potential.

As it becomes increasingly clearthat malocclusion is a predisposingfactor for certain chronic systemicdiseases27 that were likely neversuffered by our ancestors (e.g., apnea,hypertension, CVD, etc.), existingcriteria for determining orthodonticsuccess (e.g., well-aligned andstraight teeth, pathology-free and

esthetically-positioned jaw relation-ship, etc.) should also include factorsrelated to long-term systemic health(e.g., adequate posterior airwayvolume) (Figs. 8, 9, 10 & 11).

This framework holds significantimplications to currently acceptedtheories about malocclusion etiol-ogy, clinical diagnostic criteria,treatment option selections andultimate orthodontic treatmentsuccess; these and other EOM-related issues will be addressed in aproposed follow-up to this paper.

Summary & ConclusionsFor millions of years, our pre-

human and anatomically modernhuman ancestors evolved a mastica-tory apparatus (MA) that was bestadapted to foods that requiredprolonged and forceful chewing ofvaried Paleolithic-type diets (e.g.,wild whole grains, fibrous fruits andvegetables, nuts, seeds, raw andcooked meats and fish, etc.).

Constantly changing feeding envi-ronments over the several millions ofyears time-span of human evolutionare known to have been an extremechallenge to our early human ances-tors. As the various pre-human(hominid) species evolved away fromtheir common ancestor with themodern chimpanzee, an MA pheno-type that was best adapted to Pale-olithic-type diets offered the bestchance for surviving and reproducing(i.e., becoming our ancestors). Inwhat is now called the AgriculturalRevolution, also called the NeolithicRevolution, sometime around the 10-century B.C.E. (Before Common Era)in the Fertile Crescent region of theMiddle East (what is now Turkey),mankind began a gradual shift tobecoming primarily sedentary agri-culturists from having been nomadichunter-gatherer/foragers (H-G/F’s) fornearly their entire existence.

When viewed from an evolution-ary timescale perspective, the Agricul-tural Revolution represented a relativelyabrupt change in mankind’s means ofacquiring food for themselves. Thisand subsequent changes in thehuman diet have been accompaniedby an increased incidence of a myriadof chronic and non-communicablesystemic diseases (CNCD’s) likeobesity, Type 2 diabetes, CVD andsome cancers.28 Likewise, modern

38 November/December 2011 JAOS

www.orthodontics.com November/December 2011 39

Fig. 8

Fig. 10 Fig. 11

Fig. 9

Fig. 8: Retruded maxilla and mandible (pre-Biobloc). Fig. 9: Forward maxilla and mandible (post-Biobloc). Notice more protrusive profile and associated increase in posterior airwayvolume (red arrows) on the post-Biobloc images. Adapted from Hockel 2011. A significant relationship (reduction) has been demonstrated between the pre-and post-treatmentposterior airway volume and the retraction distance of lower incisors in this study. Fig. 10: Posterior airway volume before bicuspid-extraction/incisor-retraction Tx. Adapted fromWang et al, Angle Orthodontist 2011. Fig. 11: Posterior airway volume after bicuspid-extraction/incisor-retraction Tx. Adapted from Wang et al, Angle Orthodontist 2011.

Western lifestyle and foods (i.e., soft-ened/highly processed, fatty, salty,sweetened, etc.) are major causes oftwo plaque-mediated (dieto-infectous)oral maladies, dental caries and peri-odontal disease.29 These dentalCNCDs, like their systemic counter-parts, are often referred to as diseasesof civilization or Western-lifestylediseases. Although many anthropolo-gists and other scientists have

suggested that malocclusion isanother Western-lifestyle relateddisease, this view does not yet seem tobe accepted by the dental community.

As dental and other allied-healthprofessionals become better informedabout the evolutionary history of thehuman genome, and how its relativeplasticity and ability to respond toharsh and ever-changing feedingenvironments has allowed us to

survive as a species into the presentday, it should become easier tounderstand that malocclusion isindeed, a dental disease of civilization.Acknowledgements The author would like to thank Robert Perlman,MD, PhD (The University of Chicago), PhilippeHujoel, DDS, PhD (The University of Washington)and John Mew, DDS (The London School ofOrthotropics) for their editorial input and sharedinterest in the topic of Darwinian Dentistry.

Editor’s Note: Article references are avail-able upon request or for download in thedigital version at www.orthodontics.com.

Figures-websourcesFigs. 1 & 2-Palmer- www.brianpalmerdds.com/pdf/section_A.pdf

Fig. 3- Palmer- www.brianpalmerdds.com/pdf/adsm_section_b.pdf

Fig. 4- Kean- www.uic.edu/classes/osci/osci590/11_1Epidemiology.htm

Fig. 5- Sanders-http://berkeley.edu/news/media/releases/2003/06/11_idaltu.shtml

Fig. 6-AMNH- www.britannica.com/EBchecked/media/36968/Artists-reconstruction-of-a-Cro-Magnon-an-early-version-of

Fig. 7- Corruccini- www.angle.org/doi/abs/10.1043/0003-3219(1989)059%3C0061%3AOADOIE%3E2.0.CO%3B2

Figs. 8 & 9- Hockel-www.hockel.com/Orthotropics.html

Figs. 10 & 11- Wang et al-www.angle.org/doi/pdf/10.2319/011011-13.1

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9. Keene, H. 1980, “History of Dental Caries in Human Populations:The First Million Years" Symposium and Workshop on AnimalModels in Cariology, Sturbridge, Massachusets.

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Appendix

Evolutionary Medicine at Durham, UKhttp://www.dur.ac.uk/ev.med/

The Evolutionary & Medicine Review http://evmedreview.com/

NESCent catalysis meeting-Evolution of Human Teeth andJaws: Implications for Dentistry and Orthodonticshttp://www.nescent.org/science/awards_summary.php?id=309

The Ancestral Health Societyhttp://ancestryfoundation.org/AHS.html

Y Crooked Teethhttp://www.facebook.com/YCrookedTeeth

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