Complete Mitochondrial DNA Genome Sequences for Two Lineages in Coilia mystus (Clupeiformes: Engraulididae): Mitogenomic Perspective on the Phylogenetic Relationships of Genus Coilia

Ai Guo1,2,3, Jiaguang Xiao4, Binbin Shan4, Tianxiang Gao5 and Yongdong Zhou3,*

1College of Marine Sciences, Shanghai Ocean University, Shanghai 201306, China

2State Key Laboratory of Satellite Ocean Environment Dynamics, The Second Institute of Oceanography, State Oceanic Administration, Hangzhou 310012, China

3Marine Fisheries Research Institute of Zhejiang Province, Zhoushan 316021, China

4Fishery College, Ocean University of China, Qingdao 266003, China

5Fishery College, Zhejiang Ocean University, Zhoushan 316022, China

Ai Guo and Jiaguang Xiao contributed equally to this study.

ABSTRACT

To better understand the genetic diversity and phylogeny of Coilia, the complete mitochondrial genomes of two lineages in Coilia mystus were compared. They were all typical circular double stranded DNA molecule with 17075 bp in C. mystus N and 16964 bp in C. mystus S, respectively, containing the standard metazoan set of 22 transfer RNA genes, 2 ribosomal RNA genes, 13 protein-coding genes and non-coding regions. The mitogenomes of C. mystus N and C. mystus S shared the identical structural organization and gene arrangement with those of other Coilia fishes. Both lineages of C. mystus showed similar features in not only the strand-specific asymmetry of nucleotide composition, but also the codon usage of genes. Whereas a significant variation among Coilia species was detected in length of the control region, mainly caused by the variable number of tandem repeats. Phylogenetic analysis was performed based on 13 concatenated mitochondrial protein-coding genes from 8 Coilia mitochondrial genomes. The results supported that C. lindmani at first clustered with C. nasus and C. grayii which had close relationships (d=0.028), then clustered with C. mystus which exhibited obvious genetic differentiation between C. mystus N and C. mystus S (d=0.083). C. reynaldi was at the basal part of the trees, and showed obvious genetic differentiations with other Coilia species (d>0.19). Our results suggested that the north and south lineages of C. mystus could be genetically distinct as different species.

Article Information

Received 17 June 2017

Revised 24 August 2017

Accepted 02 October 2017

Available online 06 September 2018

Authors’ Contribution

AG, JX, TG and YZ conceived and designed the experiments. JX and BS performed the experiments and analyzed the data. AG, TG and YZ contributed reagents/materials. AG, JX and BS wrote the article. TG and YZ proofread the manuscript and approved the final version.

Key words

Coilia mystus, Mitochondrial genome, Genome structure, Genetic divergence, Phylogeny.

DOI: http://dx.doi.org/10.17582/journal.pjz/2018.50.6.2141.2151

* Corresponding author: zyd511@126.com

0030-9923/2018/0006-2141 $ 9.00/0

Copyright 2018 Zoological Society of Pakistan

Introduction

As an important functional eukaryotic organelle, mitochondria become highly economized and conserved (Boore, 1999). Most genes of the original endosymbiont are lost or have been transferred to the nucleus after the Genome Reductive Evolution (GRE) process (Andersson and Kurland, 1998; Khachane et al., 2007; Ghiselli et al., 2013; Kannan et al., 2014). In animals, mitochondrial DNA (mtDNA) is a compact double-stranded circular molecule that typically contains 13 protein-coding genes (PCGs), 2 ribosomal RNA genes and 22 transfer RNAs. Moreover, a large noncoding control region (CR) commonly related to the initiation of transcription and replication sequences usually presents in CRs (Boore 1999; Breton et al., 2014). The mtDNA genomes have played a significant role in the studies of population genetics and reconstruction of phylogeny due to their intrinsic properties (i.e., fast evolutionary rate, high information content, maternal inheritance and lack of recombination) (Simon et al., 2006).

Fish of the genus Coilia are small to moderate in size and primarily inhabit in coastal waters or estuaries in the Indo-West Pacific region, and some of them can tolerate low salinities in freshwater (Wongratana, 1980). Taxonomical debates and genetic divergence of Coilia, especially Coilia nasus and Coilia mystus in China, are always the hot topics of research. At present, it is commonly accepted that Coilia brachygnathus and Coilia nasus taihuensis are synonym of C. nasus (Yang et al., 2010), because small genetic divergence has been found between diadromous and freshwater ecotypes of C. nasus (Cheng and Lu, 2005; Yang et al., 2010). C. mystus which is widely distributed along the coast of China is a short distance migration fish (Whitehead, 1985). Initially, three clades of C. mystus were described as the Yangtze River group, the Minjiang River group and the Pearl River group, respectively (Cheng et al., 2008; Yan et al., 2009). Recently, the updated conclusion revealed C. mystus should be divided into two clades: C. mystus Northern populations (N) and C. mystus Southern populations (S) (Yang, 2012). The C. mystus (N) which was distributed in the north of the coast of Ningbo in China, was just the above-mentioned Yangtze River group; the C. mystus (S) which widely lived in the rest of southern coast of China consist of the Minjiang River group and the Pearl River group as previously described (Yang, 2012). Actually, there is no significant differentiation in morphological characters between C. mystus (N) and C. mystus (S), excepting on the number of gill rakers and vertebra (Yang, 2012).

In this study, we compared the complete mitogenomes of five Coilia species to explore the divergence among different species and lineages of C. mystus. Phylogenetic analysis was conducted based on the protein coding genes of the mitogenomes using the Maximum Likelihood (ML) and the Bayesian Inference (BI) methods to gain insight on the genetic diversity between two lineages of C. mystus and its phylogenetic status in genus Coilia.

 

Materials and methods

Samples collection and sequencing

Two specimens of C. mystus, namely, C. mystus Northern populations (N) and C. mystus Southern populations (S), collected from Shanghai and Daya Bay in the coast of China, respectively, were sequenced in this study. The identifications of C. mystus N and C. mystus S were based on the results of Yang (2012). Moreover, six mitochondrial genome sequences were downloaded from GenBank for genetic comparation and phylogenetic reconstruction, including Coilia grayii, Coilia lindmani, Coilia reynaldi and three specimens of C. nasus (Table II, Lavoue et al., 2010; Zhang et al., 2016, a, b; Zhao et al., 2016). Total genomic DNA was isolated from the muscle tissue by proteinase K digestion followed by the standard phenol/chloroform method (Sambrook and Russell, 2001).

The complete mitogenomes of C. mystus were amplified using a long-PCR technique (Chang et al., 1994; Miya and Nishida, 1999) and primer-walking method. Both PCR primers were designed referring to congeneric mitogenome sequence available in GenBank and implemented in Primer Premier 5.0 software (PRIMER Biosoft International). Contiguous segments overlapped by at least 50 bp to ascertain the accuracy of sequencing. Long-PCR and normal PCR reactions were performed in a TAKARA thermal cycler following the standard procedures (Cheng et al., 2012). All fragments were sequenced on ABI Prism 3730 from both strands after purification.

Sequence annotation and analysis

Sequences were edited and aligned using DNASTAR software (DNASTAR, Madison, WI, USA) with default parameters, and refined manually. Locations for protein-coding genes and rRNAs were identified by DOGMA (Wyman et al., 2004). Gene predictions were further improved by comparing DNA sequences with those of C. nasus, C. lindmani and C. reynaldi. The base composition and codon usage of the 13 protein-coding genes were analyzed with Mega 5.0 (Tamura et al., 2011). Nucleotide composition skew analysis was carried out with the formulas AT-skew = [A - T] / [A + T] and GC-skew = [G - C] / [G + C], respectively (Perna and Kocher, 1995). Most tRNA genes were identified by their proposed cloverleaf secondary structures using web-based tRNAscan-SE software (http://lowelab.ucsc.edu/tRNAscan-SE/). The remaining tRNA genes unidentified were determined by inspecting sequences for tRNA-like secondary structures and anticodons. Control region was identified by comparing with the homologous sequences. Variable number of tandem repeats (VNTRs) were detected using Tandem Repeats Finder (Benson, 1999).

Phylogenetic reconstruction

Phylogenetic analysis was performed based on 13 concatenated mitochondrial PCGs from 8 Coilia mitochondrial genomes, Engraulis japonicas (NC_003097) (Inoue et al., 2001a) and Engraulis encrasicolus (NC_009581) (Lavoue et al., 2007) were used as the out-groups. Nucleotide sequences from all 10 mitochondrial PCGs were edited and aligned using ClustalX 1.83 under default settings (Thompson et al., 1997), stop codons and gaps were removed and finally concatenated into a sequence matrix (11,391 sites in length).

The phylogenetic trees were built using two approaches including maximum-likelihood (ML) analysis by PAUP* 4.0 (Swofford, 2002) and a partitioned Bayesian inference (BI) analysis by Mrbayes 3.12 (Huelsenbeck and Ronquist, 2001). Substitution model selection was conducted by a comparison of Akaike Information Criterion (AIC) scores (Akaike, 1974) with jModelTest 2 (Darriba et al., 2012). GTR + G model was chosen as the best-fitting model for ML analyses and the node reliability was estimated after 1000 bootstrap replicates. For the Bayesian procedure, a set of optimal models was selected for different positions (GTR+I+G for the 1st and the 3rd positions, GTR+I for the 2nd position). Four Markov chains were run for 1,000,000 generations by sampling the trees every 1000 generations. After the first 2500 trees (25%) were discarded as burn-in, the 50% majority rule consensus tree and the Bayesian posterior probabilities (BPP) were estimated using the remaining 7500 sampled trees.

 

Table I.- Characteristics of the mitochondrial genomes of C. mystus N and C. mystus S.

Gene/Region	Position		Size (bp) Nucleotide (N/S)	Am ino acid	Gapb (N/S)	Codon		Str and
	From(N/S)	To (N/S)				Start	Stopa
tRNAPhe	1	69	69		0			H
12S rRNA	70	1022	953		0			H
tRNAVal	1023	1094	72		0			H
16S rRNA	1095	2786	1692		0			H
tRNALeu(UUR)	2787	2861	75		0			H
ND1	2862	3836	975	324	1	ATG	TAA	H
tRNAIle	3838	3909	72		-1			H
tRNAGln	3909	3979	71		-1			L
tRNAMet	3979	4047	69		0			H
ND2	4048	5093	1046	348	0	ATG	TA	H
tRNATrp	5094	5163	70		2			H
tRNAAla	5166	5234	69		1			L
tRNAAsn	5236	5308	73		0			L
OL	5309	5342/5341	34/33		-3			H
tRNACys	5340/5339	5405/5404	66		0			L
tRNATyr	5406/5405	5476/5475	71		1			L
CO I	5478/5477	7022/7021	1545	514	0	GTG	TAA	H
tRNASer(UCN)	7023/7022	7093/7092	71		5			L
tRNAAsp	7099/7098	7166/7165	68		11			H
CO II	7178/7177	7868/7867	691	230	0	ATG	T	H
tRNALys	7869/7868	7941/7940	73		1			H
ATPase8	7943/7942	8110/8109	168	55	-10	ATG	TAA	H
ATPase6	8101/8100	8783/8782	683	227	0	ATG	TA	H
CO III	8784/8783	9568/9567	785	261	0	ATG	TA	H
tRNAGly	9569/9568	9639/9638	71		0			H
ND3	9640/9639	9989/9988	350	116	-1	ATG	TA	H
tRNAArg	9989/9988	10057/10056	69		0			H
ND4L	10058/10057	10354/10353	297	98	-7	ATG	TAA	H
ND4	10348/10347	11728/11727	1381	460	0	ATG	T	H
tRNAHis	11729/11728	11797/11796	69		1			H
tRNASer(AGY)	11799/11798	11865/11864	67		0			H
tRNALeu(CUN)	11866/11865	11937/11936	72		0			H
ND5	11938/11937	13773/13772	1836	611	-4	ATG	TAA	H
ND6	13770/13769	14291/14290	522	173	1	ATG	TAA	L
tRNAGlu	14293/14292	14361/14360	69		5/4			L
Cyt b	14367/14365	15507/15505	1141	380	0	ATG	T	H
tRNAThr	15508/15506	15577/15575	70		-1			H
tRNAPro	15577/15575	15647/15645	71		0			L
Control region	15648/15646	17075/16964	1428/1319					H

a, TA and T represent incomplete stop codons; b, positive numbers correspond to the nucleotides separating adjacent genes, negative numbers indicate overlapping nucleotides.

 

Results and discussion

General features of the mitogenomes

The complete mitogenome sequences of C. mystus N and C. mystus S were 17, 075 bp and 16,964 bp in length (Fig. 1, Table I). Actually, C. mystus N had the longest mitogenome and the shortest was that of C. lindmani (16835 bp) in all sequenced Coilia species (Table II). Length differences were primarily the result of variation in intergenic nucleotides and the control region, predominately, variable number tandem repeats detected in control regions of all eight mitogenomes of Coilia fishes. The mitogenomes of C. mystus N and C. mystus S had the same structural organization and gene arrangement with those of other Coilia fishes (Fig. 1). Both C. mystus N and C. mystus S exhibited a clear strand-specific bias in composition, most of the genes were encoded on the heavy stand (H-strand) except for ND6 and 8 tRNAs (Gln, Ala, Asn, Cys, Tyr, Ser (UCN), Glu and Pro), which were oriented to on the light strand (L-strand). These features also could be found in other vertebrates (Lavoue et al., 2010, 2013; Li et al., 2013; Liu et al., 2017; Shan et al., 2014; Teacher et al., 2012).

The AT content of mitogenomes varied among Coilia taxa (Table II) from 56.3% (C. reynaldi) to 57.8% (C. mystus N). Overall structural compositions of Coilia, this specific bias in nucleotide composition commonly exhibited excepting the first codon positions of PCGs like other bony fishes (Mabuchi et al., 2007; Cheng et al., 2012; Xiao, 2015). For GC- / AT-skews analysis, all Coilia species were displayed a positive AT-skew from 0.069 (C. mystus N) to 0.094 (C. reynaldi), especially in rRNA genes (≥0.255), and a strong negative GC-skew from -0.257 (C. mystus N) to -0.271 (C. nasus PYH and C. grayii), especially in PCGs (≤-0.274) (Supplementary Table I).

 

Table II.- Genomic characteristics of eight Coilia mitochondrial genomes.

Species	Gen Bank acces sion	Genome		13 protein-coding genes					2 rRNA		22 tRNA		CR
		L (bp)	A+T (%)	L (bp)*	A+T (%)				L (bp)	A+T (%)	L (bp)	A+T (%)	L (bp)	A+T (%)
					All pos.	1st cod pos.	2nd cod pos.	3rd cod pos.
C. mystus S	MF36 3002	16964	57.6	11391	57.5	49.7	58.5	64.4	2645	54.0	1548	55.7	1319	67.6
C. mystus N	MF36 3003	17075	57.8	11391	57.9	49.7	58.4	65.6	2645	53.7	1548	55.0	1428	67.4
C. nasus AS	KM25 7636	16900	57.5	11391	57.5	49.5	58.5	64.6	2641	54.0	1549	55.3	1252	66.7
C. nasus NB	KM36 3243	16896	57.5	11391	57.6	49.5	58.5	64.7	2641	54.0	1549	55.3	1252	66.3
C. nasus PYH	KM2 76661	16896	57.5	11391	57.5	49.5	58.5	64.6	2641	54.1	1549	55.5	1252	67.1
C. reynaldi	NC01 4276	17064	56.3	11391	55.6	48.5	58.4	59.8	2641	53.7	1550	54.5	1419	69.2
C. grayii	KP31 7088	16851	57.2	11391	57.4	49.3	58.4	64.5	2640	53.7	1549	55.4	1208	65.2
C. lindmani	NC01 4271	16835	56.7	11391	56.9	49.0	58.4	63.2	2640	53.8	1550	55.1	1191	63.7

*excluding the stop codons; L, length; pos., position.

 

Protein-coding genes and Codon usage

All 13 protein-coding genes found in other vertebrates were also present in C. mystus N and C. mystus S as well as other Coilia species, including three subunits of the cytochrome c oxidase (COI-III), seven subunits of the NADH ubiquinone oxidoreductase complex (ND1-6, ND4L), one subunit of the ubiquinol cytochrome b oxidoreductase complex (Cyt b), and two subunit of ATP synthases (ATP6 and ATP8) (Fig. 1, Table I). Without regard to the stop codons, the length of 13 protein-coding genes were exactly same (11391 bp) among all the Coilia species (Table II). The mitogenome of C. mystus N and C. mystus S exhibited a canonical genetic code shared by most vertebrates (Inoue et al., 2001b; Ramakodi et al., 2015). An orthodox initiation codon ATG was used for all protein-coding genes except for COI starting with GTG (Table I). A diverse pattern of codon usage within stop codons consisting complete stop codon and incomplete stop codon was showed in Table I, which seems to be a common tendency in fish mitogenomes (Cheng et al., 2012; Li et al., 2013).

The mitochondrial genomes of C. mystus N and C. mystus S consisted of 3797 codons excluding stop codons. Codons for Leucine possessed the highest percentage value of 15.39% and 16.01%, which may be related to the function of chondriosome of encoding many transmembrane proteins (Gillespie et al., 2006), while those for Cysteine were the least represented with a percentage value of 0.82% and 0.81%, as observed in other Coilia fishes (Supplementary Fig. S1; Zhang, 2015). Comparing the relative synonymous codon usage (RSCU) of Coilia, it showed the similar tendency in H-strand that A-terminal was in great abundance and G-terminal was extremely destitute (Fig. 2). The underlying mechanism responsible for the strand bias has been generally interpreted as evidence of an asymmetrical directional mutation pressure associated with replication processes when one strand remains transiently in a single-stranded state, making it more vulnerable to DNA damage (Perna and Kocher, 1995).

Transfer and ribosomal RNA genes

The complete set of 22 tRNA genes which were usually found in metazoans was present in C. mystus N and C. mystus S mitogenome. In addition, 14 tRNA genes were transcribed on the H-strand, whereas other 8 tRNA genes were oriented to the L-strand (Table I). Although G-U wobbles and other atypical pairings were constantly detected, typical cloverleaf secondary structures were shown for 21 tRNA genes with the exception of tRNASer (AGY) who lacked the recognizable DHU stem found in almost all vertebrate mitogenomes (Li, 2014; Miya and Nishida, 1999; Zhang et al., 2016a). Stem mismatches were common for mitochondrial tRNA genes and were probably repaired via a post-transcriptional editing process (Lavrov et al., 2000).

The two rRNA genes which were identified in C. mystus N and C. mystus S with no length variation, a small (12S) subunit of rRNA comprising 953 bp long and a large (16S) subunit of 1692 bp, were also similar size to their counterparts in other Coilia mitogenomes (Table II). But subtle differences also existed, 16S rRNA in C. nasus and C. reynaldi were 1688 bp, 16S rRNA in C. grayii and C. lindmani were 1687 bp (Zhang, 2015).

Control region and sequence repeats

Mitochondrial control region is the major non-coding segment in the vertebrate mitogenome which is AT-rich and highly variable by a faster rate of evolution (Sbisa et al., 1997). This region frequently lead to the length variation in the whole mitogenome, whereas its control elements related to regulatory functions are known to be highly conserved (Arnason and Rand, 1992; Broughton and Dowling, 1994; Lunt et al., 1998). The control region of both C. mystus was located between the tRNAPro and tRNAPhe genes, and determined to be 1428 bp in C. mystus N and 1319 bp in C. mystus S. Actually, C. mystus N had the longest control region and the shortest was that of C. lindmani (1191 bp) in all sequenced Coilia species (Table II). The variable number tandem repeats (VNTRs) which was common in genus Coilia fishes (Zhu et al., 2008), was the main reason for different length among Coilia. The VNTRs were found always presenting after about 200bp at the start of control region and having different type among Coilia species. It is worth mentioning that it showed different type and size repeats between two linages of C. mystus, however it shared the identical repeat and times among three C. nasus (Fig. 3, Table III). The different type and size of repeat unit between two lineages of C. mystus provided strong evidence to support that the two clades may represent different species. The VNTRs were believed resulting from illegitimate elongation model (Buroker et al., 1990).

The structures of control region for Coilia species were identical except for the length change. By comparing with the recognition sites among Coilia species, three conservative domains were detected: the extended termination associated sequence domain (TAS), the central conserved sequence block domain and the conserved sequence block domain (Fig. 3). The motif-TACAT in TAS was easily found, and so was the complementary TAS (cTAS) motif-ATGTA in the extended termination associated sequence domain which may function as a recognition site for terminating synthesis of heavy strand (Clayton, 1991). Moreover, the VNTRs were in this domain. In the central conserved sequence block domain, CSB-F which distinguished the central conserved sequence block domain from the termination associated sequence domain, CSB-E which was always characterized by the GTGGG box and CSB-D were recognized. The conserved sequence block domain which was thought to be involved in positioning RNA polymerase both for transcription and for priming replication (Shadel and Clayton, 1997) was characterized by CSB1, CSB2 and CSB3.

 

Table III.- The variable number tandem repeats (VNTRs) among genus Coilia.

Species	Repeat unit	Rep eat unit (bp)	Repeat time
C. mystus S	ATATTATGCATTATATTACATATATATTATGGTATAGTAC	40	6
C. mystus N	GTACATACTATGCATTATATTACATATATTATGGTATA	38	9
C. nasus	ATATTACATATATTATGGTATAGTACATACTATGTATT	38	5
C. reynaldi	TACATATATGATATAGTACATACTATGCATTATATTACA	39	12
C. grayii	TATATTACATATATTATGGTATAGTACATACTATGTAT	38	4
C. lindmani	CATATTATGTATTATATTACATATATTATGGTATAGTA	38	3

 

Table IV.- Matrix of net average genetic distances based on 13 protein-coding genes sequences among genus Coilia.

	C. mystus S	C. mystus N	C. nasus	C. grayii	C. lindmani
C. mystus N	0.083
C. nasus	0.083	0.080
C. grayii	0.086	0.082	0.028
C. lindmani	0.095	0.093	0.051	0.055
C. reynaldi	0.195	0.194	0.195	0.197	0.198

 

Divergence of the two C. mystus mitogenomes

A sliding window analysis was used to quantify genome-wide nucleotide variability between C. mystus N and C. mystus S, contrastively, a same analysis between C. mystus N and C. mystus (a downloaded sequence from GenBank, KJ710625) was also performed (Fig. 4). The divergences between C. mystus N and C. mystus S were much higher (Fig. 4A) when compared with the nucleotide divergences between the C. mystus N and C. mystus (KJ710625) which was also collected from Yangtze estuary (<0.05, Fig. 4B). The higher divergences were showed in ND1, ND4, ND5, ND6 and Cyt b, while lower divergences were observed in tRNA or rRNA. From the nucleotide divergence analysis, it could demonstrate that there were obvious genetic divergences between north lineage and south lineage in C. mystus.

 

Phylogenetic analysis

ML and BI analyses are done with the concatenated nucleotide data containing 8 Coilia sequences and two outgroup taxa. The topological relationships of two phylogenetic analyses remained consistent, and all analyses provided high bootstrap support values for all internodes (Fig. 5). The resultant topology showed C. lindmani at first clustered with C. nasus and C. grayii which had close relationships. Then they clustered with the clade C. mystus which exhibited obvious genetic differentiation between C. mystus N and C. mystus S. C. reynaldi was at the basal part of the trees, and was showed obvious genetic differentiations with other Coilia species (Table IV). In accordance with the phylogenetic analyses, the genetic distance between C. mystus N and C. mystus S also revealed the obvious genetic differentiation (0.083), even higher than the genetic distance among C. nasus, C. grayii and C. lindmani (0.028~0.055). In conclusion, the mitogenomic data supported the deep intraspecific differentiations in C. mystus, revealing that C. mystus N and C. mystus S might be the different species.

For the first time, our study investigates phylogenetic relationships within genus Coilia based on the complete mitogenome sequences. The tree topologies obtained in the present study were identical regardless of the analytic method used, and were statistically well supported by high bootstrap and posterior probability values. Therefore, the result suggested that the north and south lineages of C. mystus could be genetically distinct as different species. The divergences of north and south lineages of C. mystus was probably on account of the drastic changes of sea level on ice age of Pleistocene (Yang, 2012; Liu et al., 2012). The C. mystus was separated in two shelters. Moreover, C. mystus was short-distance migration species (Whitehead, 1985). This life habit might have great influences on the divergences of north and south lineages of C. mystus.

 

Acknowledgments

We are grateful to Ms. Nan Zhang for her constructive suggestions on the manuscript and technical assistance and we gratefully thank Cunyin Dou, Yuting Wang and Yancui Chen for collecting the specimens. This work was supported by Special Fund for Agro-scientific Research in the Public Interest (No. 201303048 and No. 201303050) and the Scientific Startup Foundation of Zhejiang Ocean University (Q1505). We declare that there are no conflicts of interests regarding the publication of this article and authors are solely responsible for the contents and writing of the paper.

 

Supplementary material

There is supplementary material associated with this article. Access the material online at: http://dx.doi.org/10.17582/journal.pjz/2018.50.6.2141.2151

 

Statement of conflict of interest

Authors have declared no conflict of interest.

 

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