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Lepidoptera: Saturniidae) Based on Mitochondrial DNA Sequences

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Lepidoptera: Saturniidae) Based on Mitochondrial DNA Sequences Journal of Genetics (2019) 98:15 © Indian Academy of Sciences https://doi.org/10.1007/s12041-019-1072-7 RESEARCH ARTICLE Genetic diversity and phylogeny analysis of Antheraea assamensis Helfer (Lepidoptera: Saturniidae) based on mitochondrial DNA sequences MOUSUMI SAIKIA1∗ , RAMESH NATH2 and DIPALI DEVI1 1Seri-Biotech Unit, Life Sciences Division, Institute of Advanced Study in Science and Technology, Paschim Boragaon, Guwahati 781 035, India 2Department of Zoology, Dhing College, Dhing, Nagaon 782 123, India *For correspondence. E-mail: [email protected]. Received 20 June 2018; revised 1 October 2018; accepted 25 October 2018; published online 26 February 2019 Abstract. Antheraea assamensis Helfer, popularly known as Muga silkworm, the golden silk producer of northeast India is economically important and unique among the Saturniid silkworms. In this study, the genetic diversity and phylogeny of semi- domesticated and wild morphs of Muga silkworm collected from different geographical locations of northeast India were investigated based on the sequences of five mitochondrial loci, i.e. 12S rRNA, 16S rRNA, CoxI, Cytb and CR. All the five mitochondrial loci showed a strong bias towards higher ‘A’ and ‘T’ contents. Transitional substitutions were found to be more than the transversional substitutions. The rate of nucleotide substitution and average genetic divergence were found to be highest in CR sequences and lowest in 12S rRNA gene sequences among the morphs of Muga silkworm. The morphs collected from same geographical area had identical 12S rRNA, 16S rRNA, CoxI and Cytb gene sequences. Moreover, the 12S rRNA and 16S rRNA gene sequences of some semi-domesticated and wild morphs collected from different geographical locations were also found to be similar. In the phylogenetic trees generated based on the mitochondrial loci, mixing of semi-domesticated and wild morphs was observed as they shared the same group. The information generated in this study will help in formulating strategies to conserve the natural biodiversity present among these unique silkworms in northeast India. In addition, this will be useful in identifying diverse morphs of Muga silkworm, which will help in effective breeding programmes to improve its productivity. Keywords. genetic diversity; morph; Muga; phylogeny; silkworm. Introduction (Boore 1999). It encodes 37 genes and an A+T-rich region of variable length known as the control region in insects. Morphological similarities and differences have been used Mitochondrial DNA sequences have been widely used for to group and classify organisms. However, discriminating studies on population and molecular systematics of insects finer differences among morphs, strains, races, biotypes, as it has a number of specific biological properties, which breeds and populations is usually difficult due to the make mtDNA an appropriate marker for molecular bio- influence of environment on morphological characters. In diversity. Firstly, mtDNA is highly variable because of its recent years, DNA-based molecular markers are increas- high mutation rate, which can generate some signal about ingly, employed in diverse areas of biology, including population history over short time period. Secondly, it is evolution, ecology, phylogenetic studies, population genet- maternally inherited which means that the whole genome ics and population dynamics in both plant and animal behaves as a single, nonrecombining locus. These unique systems because of their abundant polymorphism and the properties allow the development of universal primers and fact that they are independent of environmental conditions easy recovery from small or degraded biological samples (Behura 2006). A number of nuclear as well as mitochon- due to its high copy number in most cells with a different drial DNA markers have profound uses (Morin et al. 2004). evolution rate in different regions of mtDNA. These struc- The animal mitochondrial DNA (mtDNA) is a circu- tural and evolutional characteristics of mtDNA sequences lar and double-stranded molecule of 14–20 kb in length make it a marker of choice for various studies. 1 15 Page 2 of 12 Mousumi Saikia et al. The northeastern region has been recognized as the In the present study, the genetic diversity and phylogeny centre of seribiodiversity in India. The Indian golden among different morphs of Muga silkworm collected from silkworm, Antheraea assamensis Helfer (Lepidoptera: Sat- various geographical locations of northeast India were urniidae) popularly known as Muga silkworm, is endemic estimated using mitochondrial 12S ribosomal RNA (12S to northeast India. Being polyphagous, this insect feeds rRNA), 16S ribosomal RNA (16S rRNA), cytochrome oxi- on 15 different host plant species. Silk produced by Muga dase subunit I (CoxI), cytochrome b (Cytb) and control silkworm is golden yellow in colour, which makes it region (CR) sequences. This information will provide effi- very attractive. This luminous golden Muga silk has now cient and effective measures for germplasm conservation secured geographical indications (GI) status, the recog- and utilization of these silkworms. nition under the intellectual property rights that it has its origin in the Assam region of the northeast India. Unlike the other Antheraea species, this species has the Materials and methods lowest chromosome number (n = 15) and ZZ/ZO sex chromosome system (Deodikar et al. 1962). Muga silk- Samples worm is known for its production of quality silk with natural golden colour, lustre, durability and resistant to After a brief survey on habitat of Muga silkworm, the UV radiation (Chowdhury 2001). It has better mechani- four morphs were collected from various geographical cal properties than other commercial silks, which impart locations of northeast India and were used in this study a wide range of use as a textile material (Devi et al. 2011). (table 1). The morphs were green (GM), blue (BM), orange The silk produced by Muga silkworm is touted as the sec- (OM) and wild (WM). They were coded as GM01, GM02, ond most expensive silk in the world after A. yamamai, GM03, GM04, GM05, GM06, BM01, BM02, OM01, and its cultivation contributes significantly to livelihoods WM01, WM02 and WM03 according to their geograph- of indigent peoples. Apart from the unique nature of ical origin (table 1). Twenty individuals per morph per the silk produced by the insect, its general morphology, locality were used for each experiment. As these silkworms behaviour and physiology are known to be different from are cultured for economic purposes throughout northeast those of other related species. Although Muga silkworm India, no specific permissions were required for the collec- is a monotypic species, it has four morphs, namely green, tions and experiments. In this study, national park or other blue and orange as semi-domesticated morphs and a wild protected area of land or sea, endangered and protected morph (Thangavellu et al. 1988). The first three morphs are species were not used. termed as semi-domesticated because during the life cycle of these morphs the larval period is completed in the trees Phenotypic characters under natural conditions and when the morphs are mature, they crawl down to the base of the tree. Further, they are The phenotypic characters like larval colour, cocoon picked up and are placed inside a room where they spin to colour, larval weight, cocoon weight, shell weight, shell form cocoons. They are kept there until the egg preparation ratio and voltinism of each sample were studied and val- stage. At the time of hatching, they are brought back to ues were expressed as mean ± standard deviation (SD), the tree again. In contrast, the whole life cycle is completed where n = 20. on the trees in case of the wild morph. Due to various reasons like overexploitation, deforestation, environmen- tal pollution, diseases etc., the population density of these Genomic DNA isolation morphs is reported to be showing a deteriorating trend. This depletion in population density results in low genetic Genomic DNA was extracted from the larval tissue of variation of this species (Goel and Krishna Rao 2004). each sample of Muga silkworm according to the stan- To develop a sustainable conservation programme, assess- dard procedure (Suzuki et al. 1972). Briefly, 1 g of larval ment of genetic variation and phylogenetic relationship tissue was ground in liquid nitrogen to a fine powder between and within species are a necessary prerequisite. and 10 mL of extraction buffer was added. The mix- Again, prior knowledge on genetic diversity among dif- ture was incubated at 37◦C for 2 h with occasional ferent populations of a single species can also be utilized swirling. The extraction buffer contained 0.1 M Tris- in breeding programmes for improving the productivity. HCl (pH 8), 0.25 M EDTA (pH 8), 0.01 M NaCl, Many studies have been carried out to assess the genetic 0.5% SDS and 100 µg/mL proteinase K. The DNA was diversity of silkworms (Kar et al. 2005; Velu et al. 2008; extracted twice with phenol:chloroform:isoamyl alcohol Chakraborty et al. 2015). Probably due to a narrow and (24:24:1) and once with chloroform. The supernatant endemic distribution of Muga silkworm, information on DNA was precipitated with 0.1 volume of 3-M sodium the genetic variation and phylogenetic relationships within acetate and two volumes of ice-cold absolute ethanol. Fol- this species is very scanty as compared with other insect lowing precipitation, the DNA was washed twice with species. 70% ethanol and dissolved in TE buffer. The RNAse Table 1. Places of collection and phenotypic characters of the morphs of Muga silkworm. Co- Larval Cocoon Morph Location ordinates Sample code colour Larval weight (g) colour Cocoon weight (g) Shell weight (g) Shell ratio (%) Voltinism ◦ Green Khanapara, 26 09 N GM01 Deep 9.20 ± 0.13 Bright 4.62 ± 0.95 0.53 ± 0.06 11.70 ± 1.88 Multivoltine ◦ Assam 91 41 E green golden brown ◦ Green Mangaldoi, 26 43 N GM02 Deep 9.40 ± 0.15 Bright 3.95 ± 0.89 0.48 ± 0.16 12.15 ± 1.21 Multivoltine ◦ Assam 92 03 E green golden brown Genetic diversity and phylogeny analysis of A.
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