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Background Helicoverpa armigera and Helicoverpa zea are major caterpillar pests of Old and New World agriculture, respectively. Both, particularly H. armigera, are extremely polyphagous, and H. armigera has developed resistance to many insecticides. Here we use comparative genomics, transcriptomics and resequencing to elucidate the genetic basis for their properties as pests.ResultsWe find that, prior to their divergence about 1.5 Mya, the H. armigera/H. zea lineage had accumulated up to more than 100 more members of specific detoxification and digestion gene families and more than 100 extra gustatory receptor genes, compared to other lepidopterans with narrower host ranges. The two genomes remain very similar in gene content and order, but H. armigera is more polymorphic overall, and H. zea has lost several detoxification genes, as well as about 50 gustatory receptor genes. It also lacks certain genes and alleles conferring insecticide resistance found in H. armigera. Non-synonymous sites in the expanded gene families above are rapidly diverging, both between paralogues and between orthologues in the two species. Whole genome transcriptomic analyses of H. armigera larvae show widely divergent responses to different host plants, including responses among many of the duplicated detoxification and digestion genes.ConclusionsThe extreme polyphagy of the two heliothines is associated with extensive amplification and neofunctionalisation of genes involved in host finding and use, coupled with versatile transcriptional responses on different hosts. H. armigera’s invasion of the Americas in recent years means that hybridisation could generate populations that are both locally adapted and insecticide resistant.Electronic supplementary materialThe online version of this article (doi:10.1186/s12915-017-0402-6) contains supplementary material, which is available to authorized users.

THE SCOPE OF PSYCHOLOGY.PSYCHOL~GY is the Science of Mental Life, both of ita phenomena and of their conditions. The bhenomena are such things as we call feelings, desires, cognitions, reasonings, decisions, and the like ; and, superficially considered, their variety and complexity is such as to leave a chaotic impression on the observer. The most natural and consequently the earliest way of unifying the material was, first, to classify it as well as might be, and, secondly, to affiliate the diverse mental mod;* thus found, upon a simple entity, the personal Soul, of which they are taken to be so many facultative manifestations. Now, for instance, the Soul manifests its faculty of Memory, now of Reasoning, now of Volition, or again its Imagination or its Appet,ite. This is the orthodox ' spiritualistic ' theory of scholasticism and of common-sense. Another and a less obvious way of unifying the chaos is to seek common elements in the divers mental facts rather than a common agent behind them, and to explain them constructively by the various forms of arrangement of these elements, as one explains houses by stones and bricks. The ' associationist' schools of Herbart in Germany, and of Hnme the Mills and Bain in Britain have thus constructed 8 psychdogy without a aod by taking discrete 'ideas,' faint or vivid, and showing how, by their cohesions, repulsions, and forms * Nothing is easier than to familiarize one's self with the mammttliun brain. Qet a sheep's head, a small saw, chisel, scalpel and forceps (all three can best be had from a surgical-instrument maker), and unravel its parts either by the aid of a human dissecting book,such a8 Holden's 'Manual of Anatomy,' or by the specific directions ad hoe given in such books as Foster and Langley's 'Practical Physiology ' (Macmillan) or Morrell'e 'Comparative Anatomy and Dissection of Mammalia' (Longmans).

The use of conventional chemical insecticides and bacterial toxins to control lepidopteran pests of global agriculture has imposed significant selection pressure leading to the rapid evolution of insecticide resistance. Transgenic crops (e.g., cotton) expressing the Bt Cry toxins are now used world wide to control these pests, including the highly polyphagous and invasive cotton bollworm Helicoverpa armigera. Since 2004, the Cry2Ab toxin has become widely used for controlling H. armigera, often used in combination with Cry1Ac to delay resistance evolution. Isolation of H. armigera and H. punctigera individuals heterozygous for Cry2Ab resistance in 2002 and 2004, respectively, allowed aspects of Cry2Ab resistance (level, fitness costs, genetic dominance, complementation tests) to be characterised in both species. However, the gene identity and genetic changes conferring this resistance were unknown, as was the detailed Cry2Ab mode of action. No cross-resistance to Cry1Ac was observed in mutant lines. Biphasic linkage analysis of a Cry2Ab-resistant H. armigera family followed by exon-primed intron-crossing (EPIC) marker mapping and candidate gene sequencing identified three independent resistance-associated INDEL mutations in an ATP-Binding Cassette (ABC) transporter gene we named HaABCA2. A deletion mutation was also identified in the H. punctigera homolog from the resistant line. All mutations truncate the ABCA2 protein. Isolation of further Cry2Ab resistance alleles in the same gene from field H. armigera populations indicates unequal resistance allele frequencies and the potential for Bt resistance evolution. Identification of the gene involved in resistance as an ABC transporter of the A subfamily adds to the body of evidence on the crucial role this gene family plays in the mode of action of the Bt Cry toxins. The structural differences between the ABCA2, and that of the C subfamily required for Cry1Ac toxicity, indicate differences in the detailed mode-of-action of the two Bt Cry toxins.