Epigenetics Mechanisms in Insects: a Review

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Epigenetics Mechanisms in Insects: a Review Int.J.Curr.Microbiol.App.Sci (2020) 9(5): 2961-2971 International Journal of Current Microbiology and Applied Sciences ISSN: 2319-7706 Volume 9 Number 5 (2020) Journal homepage: http://www.ijcmas.com Review Article https://doi.org/10.20546/ijcmas.2020.905.339 Epigenetics Mechanisms in Insects: A Review U. Pirithiraj1, R. P. Soundararajan2* and C. Gailce Leo Justin1 1Anbil Dharmalingam Agricultural College and Research Institute, Tiruchirappalli-620 027, India 2Horticultural College and Research Institute (Women), Tiruchirappalli-620 027, India *Corresponding author ABSTRACT Epigenetics is the study of heritable changes in gene expression that do not involve changes in the underlying DNA sequence (or) a change in phenotype without a change in genotype. Phenotype is K e yw or ds the observable or measurable characteristics of an organism i.e., height, behaviour, colour, shape, and size. Insects are being examined for their epigenetic phenomena and the underlying mechanism Epigenetics, Insects, behind it. Epigenetics is well studied in fruit fly, Drosophila melanogaster. Gene is segments of the DNA methylation, DNA sequence that store the information to synthesize proteins or RNAs that carry out specific Histone functions. Epigenetics is important in cellular differentiation which is responsible for the phenotypic modification and plasticity. Epigenetics is been investigated in plants and animals. On comparing other organisms RNAi insects possess a high degree of phenotypic variation. This variation is due to switch on and off mechanism of gene. Gene expression and repression in insects is regulated through epigenetic Article Info mechanisms i.e. DNA Methylation, Histone modification and Noncoding RNAs which influence the Accepted: phenotypic modification. In recent days insects are used as models in studying epigenetics. RNA 23 April 2020 interference (RNAi), also known as RNA silencing, refers to a set of molecular processes in which noncoding RNA molecules target and silence the expression of specific nucleic acids. Silencing Available Online: 10 May 2020 Dnmt3 expression in newly hatched honeybee larvae mimics the effect of royal jelly, namely, the larvae destined to become workers develop into queens with fully developed ovaries. The detailed mechanisms of epigenetics in insect were discussed in the paper. Introduction (Ahmad et al., 2012). Genetically modified mosquitoes created possibilities for The growing population and degrading controlling diseases like malaria (Catteruccia environment are two major problems at et al., 2003). international level. Satisfying the growing population‘s requirement is somewhat In 1994, the ―flavr savr‖ (first GM tomato) difficult. Although there were legal and social was commercialized (Yang et al., 2005). Bt constrains in utilizing the biotechnology cotton was the first GM crop approved for use researches in field level, these tools improve in India (Kapur et al., 2010). Almost in the agriculture and human life to a great extent entire living organism‘s gene-modification 2961 Int.J.Curr.Microbiol.App.Sci (2020) 9(5): 2961-2971 plays an important role in developing high phenotypic plasticity as evident in most transgenic organisms. In an organism genes of the insects i.e. holometabolous insects are controlled by some Non-gene having egg, larvae, pupa and adult stages all mechanisms. The expression and repression with same genome. The fruit fly, Drosophila of gene without change in respective DNA melanogaster is used as a model by the sequence occurs in all living organisms. geneticist for studying the biological Ultimately their study is called epigenetics. processes and nothing to get excited, they are Similar to genetic engineering this study leads preferred for studying epigenetics (Burggren, to a profitable/beneficial research outcome in 2017). Epigenetic inheritance deals with Intra the science of plant breeding, entomology, generational and intergenerational epigenetic pathology, nematology, biotechnology, inheritance. The former is concerned with the microbiology, agronomy, soil science etc in development process i.e. how an egg with a various branches of agriculture. single set of genetic instructions is able to develop into a multicellular organism made Epigenetics is one of the emerging branch of up of distinct tissues (Waddington, 1942). science yet to be reviewed and more research The later Heard and Martienssen (2014) is work has to be carried out for clear cut concerned with transmission of epigenetic configuration about the phenotypic plasticity. information to offspring. This paper is aimed ―Epigenetics term was coined by Conrad H. to describe and understand the basic Waddington in the year 1942‖. In view of knowledge on insect epigenetics to the Conrad Waddington, epigenetics is study of readers. epigenesist, i.e. contribution of genotypes to phenotypes during development DNA methylation (Waddington, 1942). According to Arthur Riggs and colleagues it was ―the study of Addition of methyl group to the cytosine in mitotically and/or meiotically heritable the genes is termed as ―DNA methylation‖. It changes in gene function that cannot be occurs in all three domains of life Archaea, explained by changes in DNA sequence‖. The Bacteria, and Eukarya (Klose and Bird, 2006; study of heritable changes in gene expression Suzuki and Bird, 2008). In insects DNA that occur without a change in DNA sequence methylation mostly occur in genes (Glastad et is known as Epigenetics. Without the bullock, al., 2011) and primarily target the the cart cannot be driven. There is ―gene‖ to transcription initiation site of genes (Fig. 1). study for geneticist but no ―epigene‖ for Nearly one half of DNA may be methylated epigeneticist. Epigenetic mechanisms are (Glastad et al., 2011; Bewick et al., 2018). essential in regulating the genome. Gene The regions where cytosine is followed by expression and repression is regulated by guanine are known as CpG sites or CpG three common mechanisms namely DNA islands. CpG methylation is common in methylation, histone modification, noncoding eukaryotes (Delcuve et al., 2009). In insects, RNAs occurs in all eukaryotic cells. methylation sites are predominantly restricted Additional research works in the recent years to the coding sequence of genes at CpG sites are proving the significance of epigenetics in i.e. pea aphid (Walsh et al., 2010). DNA plants, animals and as well as in insects. methylation is highly enriched in exon Many organisms are capable of developing sequences e.g. Holometabolous insects such distinct phenotypes from the same genotype as Coleoptera (Cunningham et al., 2015), (Glastad et al., 2013).Comparing all living Hymenoptera (Glastad et al., 2017) and organisms in the world insects contribute a Lepidoptera (Xiang et al., 2010; Hunt et al., 2962 Int.J.Curr.Microbiol.App.Sci (2020) 9(5): 2961-2971 2013). DNA methylation occurs in larger Application proportion of the genome e.g. Hemimetabolous insects such as Orthoptera Epigenetic mechanisms play key roles in (Wang et al., 2014), Hemiptera (Bewick et adaptation to a changing environment and al., 2019), Blattodea (Glastad et al., 2016). De have been implicated in the regulation of novo DNA methyltransferases (Dnmt3 phenotypic plasticity (Moczek and proteins) and maintenance DNA methyl Snell‐ Rood, 2008). Nemoria arizonaria transferases (Dnmt1 proteins) are the enzymes caterpillars mimic different plants depending that facilitate DNA methylation (Lyko, 2018). on their food uptake (Nijhout et al., 2009). Gene expression is inhibited by DNA Interest on the role for DNA methylation in methylation due to interference with DNA- insects surged after an experiment reduced binding transcription factors in promoter expression of dnmt3 in honey bee larvae and regions (Watt and Molloy, 1988). led to a dramatic shift from worker to queen developmental fate (Kucharski et al., 2008). Low level DNA methylation Histone modification Very low methylation rates of approximately 0.11% were observed in the genome of the The eukaryotic DNA is wrapped around silkworm moth, Bombyx mori, but all of this histone octamers, which consist of four methylation occurs in CpG islands (Xiang et different histone proteins, H2A, H2B, H3 and al., 2010). In dipteran insects dramatic loss of H4. The N-terminal tail of histone protein is DNA methylation has been observed, where post-transcriptionally modified. Histones play Dnmt1 or Dnmt3 proteins have not detected in important roles not only in regulation of gene the genome sequencing projects (Hung et al., expression but also in DNA repairing 1999; Tweedie et al., 1999; Marhold et al., mechanisms, DNA replication and 2004). CpG methylation is virtually recombination. Histones are the small basic undetectable in most developmental stages of proteins that together with DNA form D. melanogaster due to absence of Dnmts chromatin structures in the cell nucleus. (Zemach et al., 2010). Histones are modified at several different amino acid residues and with many different High level DNA methylation modifications (Lennartsson and Ekwall, 2009). Chromatin arrangement (the level of Unusually high levels of DNA methylation gene transcription) is relaxed or condensed occur in the desert locust Schistocerca according to the degree of acetylation, gregaria, which also has an unusually high methylation, phosphorylation and/or level of phenotypic plasticity, suggesting ubiquitylation of associated histones DNA methylation involvement in insect (Burggren, 2017). Replacement of canonical polyphenism
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