棘皮動物と脊椎動物の進化学的類縁性の証拠

誌名誌名弘前大学農学生命科学部学術報告 = Bulletin of the Faculty of Agricultureand Life Science, Hirosaki University

ISSNISSN 13448897

巻/号巻/号 13

掲載ページ掲載ページ p. 33-37

発行年月発行年月 2011年2月

農林水産省 農林水産技術会議事務局筑波産学連携支援センターTsukuba Business-Academia Cooperation Support Center, Agriculture, Forestry and Fisheries Research CouncilSecretariat

MINI-REVIEW

EVIDENCE OF EVOLUTIONARY CLOSE RELATIONSHIP BETWEEN ECHINODERMS AND VERTEBRATES

Norimasa MATSUOKA

Division of Molecular Evolution. Department of Biology. Faculty of Agriculture & Life Science.

Hirosaki University. Hirosaki 036-8561. Japan

(Received for publication October 19. 2010)

INTRODUCTION

33

The present author has been studying the biochemical systematics of echinoids and asteroids of echinoderms

by using some biochemical methods such as allozyme analysis. immunological assay. enzyme kinetic study. and

comparison of amino acid composition (Matsuoka. 1980. 1986. 1987. 1992; Matsuoka et al.. 1983; Matsuoka and Suzuki.

1989; Matsuoka and Hatanaka. 1991. etc.). Echinoderms are representative of sea-shore animals. Echinoderms consist

of five classes: sea-lily. sea-cucumber, starfish, sea-urchin and brittle-star (Fig. 1). The author reported the phylogeny

of four classes of echinoderms by the enzyme kinetic study of glucose-6-phospahte dehydrogenase (G6PD) (Matsuoka,

1992). Of these five echinodermes, echinoids have been widely used in various biological fields: developmental

biology, biochemistry, physiology and ecology. These five classes are distinct groups in morphological viewpoint.

The pentamery is the common body system in echinoderms. From some viewpoint. it has been considered that

echinoderms are evolutionary close related to vertebrates. In this nimi-review, I review about the molecular and non­

molecular evidence of evolutionary close relationship between echinoderms and vertebrates.

DISCUSSION

The present author has been studying the molecular phylogeny of echinoids and asteroids of echinoderms (Matsuoka,

1980; Matsuoka and Suzuki, 1989; Matsuoka and Hatanaka.l991. etc.). In parallel with biochemical systematics, I have

also been studying the population genetics of echinoderms by allozyme analysis. Echinoderms consist of five classes:

echinoids, asteroids, holosuloids, ophiuroids and crinoids. In this review, I report the molecular and non-molecular

evidence of the evolutionary close relationship between echinoderms and vertebrates. The evidence of evolutionary

close relationship between echinoderms and vertebrates is the followings: (1) They have the skeleton system. The

echinoids consist of coronal plate, the arms of asteroids consist of ambulacral plate, and our vertebarates consitst of

skeleton systems. (2) From the view point of developmental biology, they are similar to each other. They belong

to Deuterostomea. (3) The morphology of larva in echinoderms and vertebrates is similar to each other: The larva

is symmetry and it has gill. and it also has three coelomes. (4) In the comparative biochemical studies of glucose-6-

phosphate dehydrogenase (G6PD), we discovered that there exist two isozymes of G6PD in animal kingdom: one is

glucose-6-phosphate dehydrogense (G6PD) and the other hexose-6-phosphate dehydrogense (H6PD) (Matsuoka et al.

1977: Matsuoka and Hori, 1980). We investigated the distribution of the two isozymes (G6PD and H6PD) in animal

Corresponding author: Tel. +81-172-39-3590 Fax. +81-172-39- 3590 E-mail: matslloka@cc.hirosaki-ll.ac.jp

BuII.Fac.Agric.&Lifc Sci. Hirosaki Univ. No.13: 33-37. 2010

34 Matsuoka

Table 1. Comparison between G6PD and H6PD

Parameter G6PD H6PD

substrates G6P G6P,Ga16P,dG6P,Glucose

specificity

co-enzyme NAPD NADP,NAD

specificity

molecular weight 100,000 200,000

reaction to protease inactive active

distribution in cell cytosol mIcrosome

chromosome X-chromosome autosome

distribution in all animals echinoderms, vertebrates

animals

These data are quoted from Matsuoka and Hori (1980).

kingdom. As a result. we found that echinoderms and vertebrates have the two G6PD isozymes (G6PD and H6PD)

(Matsuoka and Hori. 1980). The other invertebrates except echinoderms have G6PD. but they have not H6PD. The

two isozymes are different to each other in some parameters (Table 1). G6PD has narrow substrate specificity (the

substrate is G6PD and the co-enzyme NAPD). but II6PD has broad substrate specificity: it can catalyze glucose-6-

phosphate (G6P). galactose-6-phosphate (GaI6P). deoxyglucose-6-phosphate (dG6P) and glucose. Further. H6PD also use

two co-enzymes (NADP and NAD). The two isozymes (G6PD and H6PD) have molecular weight of 100.000 and 200.000.

respectively. The chromosomes holding genes coding the two G6PD isozymes are different to each other: the gene

of G6PD is on X-chromosome. but that of H6PD on autosome. The two G6PD isozymes are homologous enzymes that

derived from common ancestor. I indicated that the two G6PD isozymes (G6PD and H6PD) are very similar to each

other in the amino acid compositions (Matsuoka et aL 1983) (Fig. 2). This similarity of amino acid composition shows

that G6PD and H6PD are homologous enzymes that derived from common ancestor. (5) Previously. we studied the

protease of the starfish. Asterias amurensis. In the study. we found that the starfish protease is trypsin-like enzyme

that homologous to vertebrate trypsin (Hori et al.. 1977). Figure 3 shows the active band pattern of starfish protease in

polyacrylamide gel electrophoresis. The starfish protease exists as inactive form in zymogen granules of pyloric caeca.

and it is activated by starfish protease as well as vertebrate trypsin. (6) The echinoids have thyroid hormone (Chino et

aL 1994; Saito et al.. 1998). The echinoids use the thyroid hormone in metamorphosis from larva to adult. The hormone

is homologous to thyroid hormone of vertebrates thyroxine. (7) In more recently. Kishimura (unpublished data) found

that the asteroids have the insulin-like hormone that reduce the blood glucose. (8) The molecular phylogenetic study

on DNA level indicated that echinoderms are closely related to vertebrates (Winnepenninckx et aL 1995)(Fig. 4). The

molecular marker used in the study is ISS ribosomal RNA. (9) Miyata (1994) demonstrated that echinoderms are

closely related to vertebrates by using multi-protein sequence analysis (Fig. 5). (10) In more recent years. Davidson

(2006) reported that the genome of echinoid (Strongylocentrotus purpuratus) is very similar to the genome of human at

DNA level. From these many evidence. I would like to propose that echinoderms are very important animal group in

evolutionary and phylogentic position in animal kingdom.

Evolutionary close relationship between echinoderms and vertebrates

~

(~' I 2

4

5 Fig. 1: Echinoderms consist of five classes: l =sea-liliy. 2=sea-c ucumber. 3=slarflsh. 4=sea-urchin. and 5=bri ttle-star.

(A) Crucian carp

Leu ~ - __ _ \ --~-.:.-............. ,

\

Val

10%

0%

.. Gly

o--oG6PD • ---eH6PD

His Arg

' ... ___ ______ ~ Asx ,

, ,

Thr

, , Ser

(8) Rat

Leu __ ---__

"'''''''" ,-: ,-----....:-::..:-'-"~

Val , ,

10·/ •

Lys

AI:---__ , -. Gly

, ,

Thr

\,Ser , \ ,

Glx

35

Asx

Fig. 2: The evidence of the similarity of amino acid sequences of G6PD and H6PD. Compar ison of amino acid compositions between G6PD and H6PD in crucian carp (A) an d ra t (E) is very si milar. T he fi gure was Cj uoted from Matsuoka et al (1983).

36 Matsuoka

+

Fig. 3: The band pattern of protease of the starfish Asterias amurensis in polyacrylamide gel electrophoresis. The two active bands are white bands in blue-background. The stain of bands was performed in amide black.

65

100

100

Chordata (Xla) Deuterostomes Chordata (Hsa) ]

Hemichordata (Sko) Echinodermata (Spu) Arthropoda (Asa) Arthropoda (Tmo) Arthropoda (Eca) Nematomorpha (Gaq) Priapullda (Pca) Mollusca (Lka) Mollusca (Aja) Mollusca (Pma) Annelida (Efe) Annelida (Lco) Acanthocephala (Mmo) Rotifera (Bpi) Gastrotricha (Lsq) Platyhelminthes (Ovi) Nematoda (Pty) Nematoda (Cel) Nemadota (Hpl) Nematoda (Nba)

Protostomes

Cnidaria (Aku) O· I bl Cnidaria (Asu) J

IpO asts Ctenophora (Mle) Porifera (Sci) Fungi (Sce)

Fig. 4: Molecular Phylogenetic tree based on DNA sequence by Winnepenninckx et al. (1995). This tree shows the close relationship between echinoderms and vertebrates.

Evolutionary close relationship between echinoderms and vertebrates 37

Mollusca

Arthropoda

Echinodermata

Vertebrata

Fig. 5: Molecular phylogenetic tree based on multi-protein seqence analysis by Miyata (1994). This tree shows the close relationship between echinoderms and vertebrates.

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