<?xml version="1.0" encoding="UTF-8"?>
<TEI xmlns="http://www.tei-c.org/ns/1.0" xml:lang="en">
  <teiHeader>
    <fileDesc>
      <titleStmt>
        <title>GENETICS OF NATURAL POPULATIONS. XIX. ORIGIN OF HETEROSIS THROUGH NATURAL SELECTION IN POPULATIONS OF DROSOPHILA PSEUDOOBSCURA</title>
        <author>Theodosius Dobzhansky</author>
      </titleStmt>
      <publicationStmt>
        <p>Tag layer of the DHSS Meta-Tagging Project. The article text is reproduced from the
           staged copy named in sourceDesc and is not altered; the tags are held separately in
           standOff so that overlapping and nested spans are preserved.</p>
      </publicationStmt>
      <sourceDesc>
        <bibl>
          <title>GENETICS OF NATURAL POPULATIONS. XIX. ORIGIN OF HETEROSIS THROUGH NATURAL SELECTION IN POPULATIONS OF DROSOPHILA PSEUDOOBSCURA</title>
          <date>1950</date>
          <publisher>Genetics</publisher>
          <idno type="DOI">10.1093/genetics/35.3.288</idno>
          
          <idno type="record">dobzhansky1950-genetics-of-natural-populations-xi</idno>
          <note type="staged-file">dobzhansky1950-genetics-of-natural-populations-xi.txt, the project's staged copy</note>
          <note type="text-sha1">35751fcbd3ca05da51bd9fb4158590c8810961e6</note>
          <note type="rights">open-access</note>
        </bibl>
      </sourceDesc>
    </fileDesc>
    <profileDesc>
      <textClass><keywords><term>population-genetics</term></keywords></textClass>
      <particDesc>
        <person xml:id="paper"><p>the paper's own words</p></person>
        <person xml:id="tagger"><p>the tag layer; a sentence written by the tagger about the
          paper, always anchored to a quotation but not itself a quotation</p></person>
      </particDesc>
    </profileDesc>
    <revisionDesc>
      <change when="2026-10-06"
              who="build_paper_edition.py">0 spans located by searching the stored text for the quote's exact words, allowing any whitespace between them</change>
    </revisionDesc>
  </teiHeader>
  <standOff>
    <spanGrp type="tags">
      <span xml:id="t0" type="claim_type" ana="#claim_type" resp="#paper">
        <label>mechanism</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t1" type="phenomenon" ana="#phenomenon" resp="#paper">
        <label>heterosis in chromosomal inversion heterozygotes</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t2" type="method" ana="#method" resp="#paper">
        <label>experiment</label>
        <term>experiment</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t3" type="mechanisms" ana="#mechanisms[0]" resp="#tagger">
        <label>mechanisms</label>
        <term>Natural selection coadapts polygene complexes only within a population, so chromosomes brought together from populations that never interbreed have not been mutually adjusted and their heterozygotes lack heterosis.</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t4" type="mechanisms" ana="#mechanisms[1]" resp="#tagger">
        <label>mechanisms</label>
        <term>Heterosis arises from interaction between polygene complexes carried by differently arranged chromosomes, which the inversions protect from being broken up by crossing over in the heterozygote.</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t5" type="mechanisms" ana="#mechanisms[2]" resp="#tagger">
        <label>mechanisms</label>
        <term>A second, distinct kind of heterosis arises when deleterious recessive mutants accumulated in the population are sheltered by their normal dominant alleles in the heterozygote.</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t6" type="mechanisms" ana="#mechanisms[3]" resp="#tagger">
        <label>mechanisms</label>
        <term>Viability and adaptive value diverge because larval survival is only one of several components (adult longevity, fecundity, sexual activity) that jointly determine net fitness.</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t7" type="limitations" ana="#limitations[0]" resp="#tagger">
        <label>The cages were not run long enough for the inferior chromosome type to be eliminated, so the predicted outcome was not observed to completion.</label>
        <note>what this limits: the elimination prediction specifically: THE CH CHROMOSOMES WERE RARE IN CAGES NOS. 48 AND 49, BUT MUCH LONGER OBSERVATION WOULD BE NEEDED TO SEE THEM ELIMINATED COMPLETELY</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t8" type="limitations" ana="#limitations[1]" resp="#tagger">
        <label>In the California-Mexico cage No. 55 the viability differences among genotypes did not reach conventional significance.</label>
        <note>what this limits: the viability comparison in cage No. 55: THE DIFFERENCES ARE, HOWEVER, MOSTLY BELOW THE CONVENTIONAL LEVEL OF STATISTICAL SIGNIFICANCE</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t9" type="limitations" ana="#limitations[2]" resp="#tagger" cert="1.0">
        <label>The paper opens its discussion by saying the term names several scarcely related phenomena that the field treats as one.</label>
        <note>what this limits: any result reported as heterosis or hybrid vigor: the paper states these terms Quotation withheld, so two such results need not be about the same thing</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t10" type="limitations" ana="#limitations[3]" resp="#tagger" cert="1.0">
        <label>Fitness measured at one developmental stage can point the opposite way from fitness overall, and the paper gives its own case.</label>
        <note>what this limits: a fitness or survival figure from a single developmental stage: adaptive value is stated not to be proportional to survival value at all stages, and here some types with higher egg-to-adult mortality were nevertheless adaptively superior</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t11" type="limitations" ana="#limitations[4]" resp="#tagger" cert="1.0">
        <label>The effect is a property of chromosomes that evolved together, so the same gene-arrangement label does not carry the same behaviour between localities.</label>
        <note>what this limits: a heterosis result carried from one population to another: polygene complexes from geographically distant localities have not undergone coadaptation, so the same chromosomal type does not behave the same way in a different population</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t12" type="controls" ana="#controls[0]" resp="#tagger" cert="0.85">
        <label>inversion heterozygotes compared against inversion homozygotes, with the paper noting that the heterozygotes were also heterozygous for geographical gene complexes</label>
        <note>eliminates: very little on its own - and the paper says so, which is why this is worth recording: it names the confound its own comparison does not remove · outcome (stated by the tagger, no quotation): partial</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t13" type="concepts" ana="#concepts[0]" resp="#paper">
        <label>heterosis</label>
        <term>heterosis</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t14" type="concepts" ana="#concepts[1]" resp="#paper">
        <label>euheterosis</label>
        <term>euheterosis</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t15" type="concepts" ana="#concepts[2]" resp="#paper">
        <label>luxuriance</label>
        <term>luxuriance</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t16" type="concepts" ana="#concepts[3]" resp="#paper">
        <label>adaptive value</label>
        <term>adaptive value</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t17" type="concepts" ana="#concepts[4]" resp="#paper">
        <label>polygene complexes</label>
        <term>polygene complexes</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t18" type="concepts" ana="#concepts[5]" resp="#paper">
        <label>population cages</label>
        <term>population cages</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t19" type="senses" ana="#senses[0]" resp="#tagger" cert="1.0">
        <label>heterosis - KIND ONE, AN ACCIDENT OF MUTATION: superiority arising because deleterious recessive mutants are hidden by their dominant alleles, which is a by-product of the mutation process</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t20" type="senses" ana="#senses[1]" resp="#tagger" cert="1.0">
        <label>heterosis - KIND TWO, A PRODUCT OF SELECTION: specific interaction effects from complexes of linked polygenes, called overdominance, engendered by natural selection as adaptation</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t21" type="senses" ana="#senses[2]" resp="#tagger" cert="1.0">
        <label>adaptive value - AS FREQUENCY CHANGE IN A CAGE: not a physiological measurement but the observed change in the relative frequencies of two types over time in a population cage</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t22" type="claim_relations" ana="#claim_relations[0]" resp="#tagger">
        <label>builds-on → Wright and Dobzhansky (1946) and Dobzhansky (1947a, b) population-cage results on within-population heterosis</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t23" type="concept_edges" ana="#concept_edges[0]" resp="#tagger">
        <label>polygene complexes —causes→ heterosis</label>
        <term>polygene complexes</term>
        <term>heterosis</term>
        <note>strength of the relation: 1</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t24" type="concept_edges" ana="#concept_edges[1]" resp="#tagger">
        <label>euheterosis —distinct-from→ luxuriance</label>
        <term>euheterosis</term>
        <term>luxuriance</term>
        <note>strength of the relation: 1</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t25" type="concept_edges" ana="#concept_edges[2]" resp="#tagger">
        <label>EVIDENCE CORRECTED (RE-GROUNDED) · adaptive value —operationalised-as→ population cages</label>
        <term>adaptive value</term>
        <term>population cages</term>
        <note>EVIDENCE CORRECTED: RE-GROUNDED. Same sentence, CARRIED TO ITS END. The old span stopped at &#x27;observing the…&#x27; and lost the apparatus; the full sentence finishes &#x27;…breeding in POPULATION CAGES…&#x27;, which is the operationalisation the edge records. The broken word &#x27;popula tions&#x27; is the staged text&#x27;s own and is quoted as it stands. — strength of the relation: 1
Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but adaptive values are DETERMINED by observing frequencies in population cages; operationalised-as is loose.</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t26" type="content_tags" ana="#content_tags.statistics[0]" resp="#tagger">
        <label>chi-squared</label>
        <term>chi-squared</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t27" type="content_tags" ana="#content_tags.statistics[1]" resp="#tagger">
        <label>degrees of freedom</label>
        <term>degrees of freedom</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t28" type="content_tags" ana="#content_tags.statistics[2]" resp="#tagger">
        <label>significance level</label>
        <term>significance level</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t29" type="content_tags" ana="#content_tags.findings[0]" resp="#tagger">
        <label>Inversion heterozygotes built from two third chromosomes of the same population show heterosis, but heterozygotes built from chromosomes of geographically remote populations generally do not.</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t30" type="content_tags" ana="#content_tags.findings[1]" resp="#tagger">
        <label>In every experiment one homozygote was sharply less viable than the heterozygote (quotients 0.17 to 0.76).</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t31" type="content_tags" ana="#content_tags.findings[2]" resp="#tagger">
        <label>Heterozygosity for same-population ST and AR third chromosomes raises viability even on a geographically mixed background of the other chromosomes.</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t32" type="content_tags" ana="#content_tags.findings[3]" resp="#tagger">
        <label>In both cages the adaptive values were ST/AR &gt; ST/ST &gt; AR/AR, the reverse of the viability relations in cage 46.</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t33" type="content_tags" ana="#content_tags.findings[4]" resp="#tagger">
        <label>By March 1948 Arrowhead had risen to 91 percent and Chiricahua had fallen to 9 percent.</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t34" type="content_tags" ana="#content_tags.findings[5]" resp="#tagger">
        <label>In cage 56 the Pinon Arrowhead homozygote was significantly more viable than the heterozygote, and the Mexican Chiricahua homozygote equally or slightly more viable.</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t35" type="content_tags" ana="#content_tags.findings[6]" resp="#tagger">
        <label>A type&#x27;s adaptive value (net fitness) is not necessarily proportional to its survival at every developmental stage.</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t36" type="content_tags" ana="#content_tags.definitions[1]" resp="#tagger">
        <label>coadaptation</label>
        <term>coadaptation</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t37" type="content_tags" ana="#content_tags.definitions[2]" resp="#tagger">
        <label>viability quotient</label>
        <term>viability quotient</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t38" type="content_tags" ana="#content_tags.quantities[0]" resp="#tagger">
        <label>0.17 to 0.76 — Range of viability quotients of the inferior homozygote relative to the heterozygote across the five geographically mixed cages</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t39" type="content_tags" ana="#content_tags.quantities[1]" resp="#tagger">
        <label>chi-squares 6.63 to 16.05 — Range of chi-squared values testing the inferiority of the poorly viable homozygote against the heterozygote, on the pooled sexes</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t40" type="content_tags" ana="#content_tags.quantities[2]" resp="#tagger">
        <label>91 percent ... 9 percent — Frequencies of Mather Arrowhead and Pinon Chiricahua chromosomes in cage 45 by March 1948, from an equal start in February 1947</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t41" type="content_tags" ana="#content_tags.quantities[3]" resp="#tagger">
        <label>79 percent ... 53 percent — Equilibrium frequency of Arrowhead reached in pure Pinon Flats and in pure Mather populations, against which the runaway hybrid cages are compared</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t42" type="content_tags" ana="#content_tags.quantities[4]" resp="#tagger">
        <label>1,217 — Number of AR/CH heterozygote flies introduced into population cage No. 45 at the start of the experiment</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t43" type="content_tags" ana="#content_tags.quantities[5]" resp="#tagger">
        <label>300 miles — Distance between the two Californian source localities, Mather and Pinon Flats</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t44" type="content_tags" ana="#content_tags.quantities[6]" resp="#tagger">
        <label>31:32 — Probability that both chromosomal types of a heterozygous parent are detected when six larvae are examined, implying one thirty-second of heterozygotes are misclassified as homozygotes</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
    </spanGrp>
  </standOff>
  <text>
    <body>
      <p><note type="rights">The full text of this article is not redistributable, so it is
      not included here. The evidence sentences are withheld with it; the tag layer and its
      grounding status remain. The article itself is at https://doi.org/10.1093/genetics/35.3.288.</note></p>
    </body>
  </text>
</TEI>
