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Going on an Antedate Evolutionary Anthropology 16:204–209 (2007) CROTCHETS & QUIDDITIES Going on an Antedate A strange history of imperfect perfect proportions. KENNETH M. WEISS AND JEFFREY A. KURLAND In the mid-1700s, the English lan- tion from which leads to the changes explained the inheritance patterns of guage was rather out of control. In that define evolution. However, since quantitative traits like skin color. the absence of a satisfactory refer- it will be on the test, many students Mendel’s ideas were different. He ence, word meanings were unclear. (even those who are awake) see HW was trained to think that organic na- Even Shakespeare had already only as an algebraic threat to their ture follows the same laws as does become somewhat obscure to most grade-point average. As one student the physical world of fundamental readers. So the intellectual-about- griped, ‘‘Let me get this straight. particles.4–6 Inheritance must follow town, Samuel Johnson, convinced When nothing happens . nothing mathematical rules. He used hybrid- investors that an English dictionary changes? Duh.’’ ization experiments to deduce those based on improved principles would Over decades of textbooks, HW rules from the relative frequency of be commercially successful.1–3 Exist- appears to be taught largely because alternative traits such as green ver- ing dictionaries were based on ‘‘a it always has been taught. It’s part of sus yellow peas. He succeeded genealogy of sentiments,’’ as author our pedagogic heritage, but its mean- because he deliberately chose traits had copied author over time. Instead, ing and rationale have become hid- known to breed true in different Johnson argued for an approach that den in the dusty volumes of past strains of peas, and he showed that reflected ‘‘intellectual history,’’ a term journal articles. It seems unthinkable the rules were general by repeating he appears to have coined (Fig. 1). to omit it. But why? To see that, we his experiments and testing seven Appearing in 1755, ‘‘Johnson’s’’ be- have to go back to the origin of mod- unrelated traits. came the standard dictionary for ern genetics. Mendel avoided traits with blurry about a century. He showed a word’s intermediates, but he did explain a evolving meaning by using historical fact that also puzzled Darwin and examples of its usage. FIRST PRINCIPLES FIRST Galton. A trait could disappear in In science as well as literature, When Gregor Mendel did his fa- one generation, then reappear in the meaning can become blurred over mous experiments with peas, the na- next. If yellow and green peas were time. Perhaps even more problematic ture of inheritance was unknown. crossed, for example, all the off- and harder to trace than word defini- Following an ancient idea, Darwin spring plants bore yellow peas. But if tions are changes in concepts. An thought that each organ transmitted he then crossed those plants, the interesting example is the Hardy- a tiny miniature of itself to the green trait would reappear in the Weinberg principle (HW). After gonads and that the ‘‘gemmules’’ next generation and—a vital clue— teaching the essentials of Darwinian from each parent blended to form the proportions of the two types evolution, Mendelian inheritance, the offspring. It was quickly shown were predictable and repeatable. In molecular biology, and speciation, that blending didn’t work, but Dar- particular, in the offspring cross he students are routinely introduced to win’s cousin Francis Galton ad- found his famous 3:1 ratio that HW. We teach it as a stable genetic vanced a modified theory of Ances- always favored one trait. He inferred baseline, the HW equilibrium, devia- tral Heredity: You receive a set of that a plant inherited nonblending heritable units from your parents. factors from each parent, and was The fertilized egg uses up half of that equally likely to transmit either to its Ken Weiss and Jeff Kurland are in the material to construct you as an orga- offspring. In modern terms, if a yel- Department of Anthropology at Penn nism; the remaining half is saved to low-producing allele (variant state) A State University. be doled out to your children, who is dominant over the green a, the get half their material from you and original AA 3 aa (yellow 3 green) half from your spouse. Thus, your cross produces only yellow Aa off- direct contribution is cut by half in spring. Listing the maternally VC 2007 Wiley-Liss, Inc. each succeeding generation, gradu- derived allele first, when you cross DOI 10.1002/evan20151 Published online in Wiley InterScience ally petering out until there is hardly these with each other, you get equal (www.interscience.wiley.com). any left. This ‘‘biometric’’ theory numbers of AA, aA, Aa, and aa CROTCHETS & QUIDDITIES Antedating Ourselves 205 Figure 1. Some defining wisdom from Samuel Johnson’s Preface, 1755.1 offspring. Since only the aas are In 1902, the English statistician stops, the then-current ratio of domi- green, that is Mendel’s famous 3:1 George Udny Yule asked what would nants to recessives, whatever it is, ratio. happen to Mendelian proportions if will remain indefinitely. So depend- you hybridized two strains (AA 3 ing on when selective breeding stops, aa) and let their descendants ran- there are any number of possible sta- TODAY’S STUDENTS NOT THE domly breed thereafter instead of ble proportions. Independently, in setting up controlled breeding for England, Karl Pearson proved that ONLY ONES CONFUSED each generation. Yule correctly the 3:1 ratio is stable in a random- Mendel’s work was recognized in inferred that under Mendel’s rules mating population. But, like Yule, he 1900, and ‘‘gene’’ was coined by the the classic 3:1 ratio would persist in started with equal allele frequencies Danish botanist Wilhelm Johannssen every future generation. Since the and never generalized these results in 1909 to refer to his inherited par- founders in Yule’s thought-experi- to all possible frequencies. The great ticles, whatever they are. As Samuel ment were AA and aa, hence the al- statistician should have done that, Johnson might find amusing, so lele frequencies were 0.5, this evolu- but he didn’t. many new functional aspects of the tionary result was a not terribly sur- By 1908, confusion was rampant. genome have been found that the prising extension of Mendel’s own A well-known Mendelian, Reginald definition has morphed to its latest experiments. But Yule then asked Punnett, presented a paper on Men- hardly-helpful incarnation of ‘‘a what would happen if only the domi- delism in humans, citing eye color union of genomic sequences encod- nant 75% of the population, rather and brachydactyly (short fingers) as ing a coherent set of potentially over- than everybody, were allowed to examples, which puzzled Yule. In a lapping functional products.’’7 breed? This would be the kind of ar- random-mating population like Eng- When we teach old, established, tificial selection that agricultural land, why didn’t Punnett find a 3:1 everybody-knows-that concepts, it’s breeders had done for centuries, and ratio of brown to blue eyes? Now it easy to forget that a century ago Darwin used as a model for evolu- was Punnett’s turn to be puzzled, these were new ideas that solved real tion by natural selection. Yule due to his inaccurate reading of scientific conundrums, and in ways wrongly calculated that in a few gen- some comments published by Yule. that were not always evident at the erations the dominants would Why did Yule think that Punnett time. approach a limiting proportion of thought that the population should Mendel’s 3:1 ratio refers to a single 85.355339%. This seemed to show increasingly be brown-eyed and bra- generation of peas from a breeding that Galton’s theory was right: The chydactylous? After all, whether experiment. But if this is a general wimpy recessive trait may brown eyes or short fingers were law of inheritance, what would hap- stay around, but diluted. But Mendel increasing or not, their inheritance pen over many generations—on the was right, too, because the recessive pattern was clearly Mendelian! evolutionary scale? This proved to be keeps reappearing. Yule concluded By this time, the relationship a surprisingly confusing question in that Mendel’s principles were a spe- among variation, heredity, breeding, which evolution, semantics, and the cial case of ancestral heredity for a selection, and evolution had been myopic eye of preconception entered dominant trait. mixed into a heady brew of concep- the drama. The effect of a genotype However, W. E. Castle, the leading tual confusion of heroic proportions. on producing a trait was confused American defender of Mendel’s Four people, Yule, Castle, Pearson, with its effect on the competitive fit- theory, was delighted to find a mis- and Punnett (Fig. 2) had independ- ness of the trait. Perhaps, in part, take in Yule’s calculation; a realiza- ently seen what the problem was, but Mendel’s word ‘‘dominirende,’’ or tion showed the failure of Galton’s each had solved only part of it. But ‘‘dominating,’’ suggested a trait’s abil- theory. If selection consistently elimi- what has all this to do with Hardy ity to bully its way to selective vic- nated the recessive every generation, and Weinberg, whoever they were? tory.4,8 Only later did the translation but all possible matings among the become ‘‘dominant’’ and thereby lose remaining dominants are calculated, any such evolutionary connotation. the dominant will very, very, very HARDY, WEINBERG, AND THEIR Johnson would be pleased. Two slowly approach 100%. There is no SILENT PARTNERS other concepts were confused, the stable proportion of recessives with descriptive word ‘‘proportion’’ and constant selection against them.
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