Letter | Published:

The industrial melanism mutation in British peppered moths is a transposable element

Nature volume 534, pages 102105 (02 June 2016) | Download Citation


Discovering the mutational events that fuel adaptation to environmental change remains an important challenge for evolutionary biology. The classroom example of a visible evolutionary response is industrial melanism in the peppered moth (Biston betularia): the replacement, during the Industrial Revolution, of the common pale typica form by a previously unknown black (carbonaria) form, driven by the interaction between bird predation and coal pollution1. The carbonaria locus has been coarsely localized to a 200-kilobase region, but the specific identity and nature of the sequence difference controlling the carbonariatypica polymorphism, and the gene it influences, are unknown2. Here we show that the mutation event giving rise to industrial melanism in Britain was the insertion of a large, tandemly repeated, transposable element into the first intron of the gene cortex. Statistical inference based on the distribution of recombined carbonaria haplotypes indicates that this transposition event occurred around 1819, consistent with the historical record. We have begun to dissect the mode of action of the carbonaria transposable element by showing that it increases the abundance of a cortex transcript, the protein product of which plays an important role in cell-cycle regulation, during early wing disc development. Our findings fill a substantial knowledge gap in the iconic example of microevolutionary change, adding a further layer of insight into the mechanism of adaptation in response to natural selection. The discovery that the mutation itself is a transposable element will stimulate further debate about the importance of ‘jumping genes’ as a source of major phenotypic novelty3.

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Primary accessions

Data deposits

The typica 1 haplotype (b–d interval) reference sequence has been deposited in GenBank under accession number KT182637; The B. betularia whole genome sequence has been deposited in the NCBI SRA database under accession number SRX1060178; the cortex splice variants have been deposited in GenBank under accession numbers KT235895KT235906; Rps3A has been deposited in GenBank under accession number JF811439; α-spec has been deposited in GenBank under accession number KT182638.


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The University of Liverpool Centre for Genomic Research (M. Hughes, C. Bourne, R. Eccles, C. Hertz-Fowler and J. Kenny) performed next-generation sequencing and Fragment Analyzer measurements. L. Cook directed us to historical data sources. C. Bergman advised on transposon detection. Population genetics simulations were performed on the University of Liverpool Advanced Research Computing Condor service. This work was supported by Natural Environment Research Council grants NE/H024352/1 and NE/J022993/1.

Author information

Author notes

    • Arjen E. van’t Hof
    •  & Pascal Campagne

    These authors contributed equally to this work.


  1. Institute of Integrative Biology, University of Liverpool, Biosciences Building, Crown Street, Liverpool L69 7ZB, UK

    • Arjen E. van’t Hof
    • , Pascal Campagne
    • , Daniel J. Rigden
    • , Carl J. Yung
    • , Jessica Lingley
    • , Neil Hall
    • , Alistair C. Darby
    •  & Ilik J. Saccheri
  2. Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Hinxton, Cambridgeshire CB10 1SA, UK

    • Michael A. Quail


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I.J.S., A.E.v.H. and P.C. designed the study and wrote the paper; P.C., A.E.v.H. and D.J.R. produced the figures; A.E.v.H. directed molecular biology experiments; A.E.v.H., C.J.Y. and J.L. conducted molecular biology experiments; A.E.v.H. constructed the BAC and fosmid tilepaths; A.E.v.H. and A.C.D. assembled, finished and annotated sequences; P.C. analysed population genetic and gene expression data; I.J.S. collected the wild sample; I.J.S. and C.J.Y. reared the samples and performed dissections; D.J.R. and A.E.v.H. built the cortex tree; D.J.R. modelled the cortex structure; M.A.Q. constructed the fosmid library; and A.C.D. and N.H. advised on the design of sequencing strategies.

Competing interests

The authors declare no competing financial interests.

Corresponding author

Correspondence to Ilik J. Saccheri.

Extended data

Supplementary information

PDF files

  1. 1.

    Supplementary Information

    This file contains Supplementary Methods, Supplementary Figures 1-2 and Supplementary References.

Text files

  1. 1.

    Supplementary Data

    This file contains the sequence alignment of the carbonaria and three typica haplotypes spanning the ‘b-d’ region (illustrated in Extended Data Figure 1).

  2. 2.

    Supplementary Data

    This file contains full-length sequence alignment in aligned FASTA format of cortex proteins and selected homologues. Incompleteness of some sequences at the N-terminus and some uncertainty regarding translation start sites have no impact on the phylogenetic tree since it was calculated using only the propeller domain (see Extended Data Figure 9a).

Excel files

  1. 1.

    Supplementary Table 1

    This table shows polymorphisms in the carbonaria candidate region.

  2. 2.

    Supplementary Table 2

    This table contains polymorphisms in the locus b-d region.

  3. 3.

    Supplementary Table 3

    This table contains Carbonaria morph frequencies in the Manchester area.

  4. 4.

    Supplementary Table 4

    This table contains PCR primers for cortex, control genes and candidate genes.

  5. 5.

    Supplementary Table 5

    This table contains sources, including accession numbers, for cortex and Fizzy family sequences.

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