Comparison of the Spawning Characteristics of the Yellow Striped Flounder Pseudopleuronectes herzensteini in the Coastal Waters off Gangwon and Gyeongbuk, Korea

한수지 54(3), 318-328, 2021

318

Copyright © 2021 The Korean Society of Fisheries and Aquatic Science pISSN:0374-8111, eISSN:2287-8815 Korean J Fish Aquat Sci 54(3),318-328,2021

Original Article

서 론

참가자미

(Pseudopleuronectes herzensteini)

^{는 황해와 보하} 이만

,

우리나라동해

,

일본의혼슈

,

북해도를거쳐쿠릴제도까 지분포하는냉수성저서어류이다

(FishBase, 2021).

최근까지 참가자미의학명은

Pleuronectes herzensteini

^{로통용되었으나}

,

본연구에서는

Ji et al. (2016)

의계통분류학적연구결과에따라

Pseudopleuronectes herzensteini

^{로표기하였다}

.

동해안가자미 류의어획량은

1990

년대평균

5,541

톤

, 2000

년대평균

8,854

톤

, 2010

년대평균

9,605

톤으로지난

30

년간증감을반복하며 증가하는추세이다

(Fig. 1) (KOSIS, 2021).

^{동해안가자미류는} 주로저인망

,

트롤

,

자망등으로어획되는어종으로상업적으로 그가치가크다

.

하지만위판장에서위판시어종을분류하지않

고

‘

^가자미류

’

로통합하여어획량을집계하고있으며

, 2015

^년 부터가자미류중기름가자미만따로분류하여어획통계를집 계하고있다

.

이러한이유로참가자미의정확한어획량을파악 하기에는 어려운실정이다

.

정부에서는지속적으로참가자미 의자원을이용하기위한방안으로포획금지체장을설정하여 관리하고있다

.

^{수산자원관리법시행령에따르면기존}

12 cm

였던포획금지체장을

2021

년부터

3

년간

17 cm

로확대실시할

예정이며

,

이후에는

20 cm

로강화할방침이다

.

포획금지체장

을설정하고검토하기위해서는대상어종의산란생태연구에 대한기초자료가필수적으로확보되어야한다

.

참가자미에관 한국내연구동향을살펴보면

,

^{난발생과정과자어의형태발달} 연구

(Han and Kim, 1999),

난발생에미치는수온과염분의영 향연구

(Lee et al., 1997),

연령과성장연구

(Choi et al., 1986;

동해안에 서식하는 참가자미(Pseudopleuronectes herzensteini)의 해역 별 산란특성 비교

김소라·이수정 ¹ ·양재형 ¹ *

국립수산과학원 연근해자원과, ¹국립수산과학원 동해수산연구소 자원환경과

Comparison of the Spawning Characteristics of the Yellow Striped Flounder Pseudopleuronectes herzensteini in the Coastal Waters off Gangwon and Gyeongbuk, Korea

So Ra Kim, Soo Jeong Lee

¹

and Jae Hyeong Yang

¹

*

Researcher, Coastal Water Fisheries Resources Research Division, National Institute of Fisheries Science, Busan 46083, Korea

1Researcher, Fisheries Resources and Environment Research Division, East Sea Fisheries Research Institute, National Institute of Fisheries Science, Gangneung 25435, Korea

We investigated the maturity and spawning of the brown sole Pseudopleuronectes herzensteini , in the coastal waters of the East Sea off Gangwon and Gyeongbuk, South Korea, using samples collected by gill net and longline fishery from January 2018 to December 2019. We analyzed oocyte development, monthly maturity stage, gonadosomatic index (GSI) and total length at maturity. Histological analysis of ovarian development in P. herzensteini revealed that the development was group-synchronous. The spawning peak in Gangwon and Gyeongbuk occurred from April to May and from February to March, respectively. Thus, the spawning season in Gangwon was 1-2 months later than that in Gyeongbuk. The length at 50% maturity for females was estimated as 24.2 cm and 19.0 cm in Gangwon and Gyeongbuk, respectively. Similarly, the length at 50% maturity for males was estimated as 19.8 cm and 16.5 cm in Gangwon and Gyeongbuk, respectively.

Keywords: Maturity, Spawning, Yellow striped flounder, Flatfish, Pseudopleuronectes herzensteini

*Corresponding author: Tel: +82. 33. 660. 8522 Fax: +82. 33. 661. 5813 E-mail address: [email protected]

This is an Open Access article distributed under the terms of the Creative Commons Attribution Non-Commercial Licens (http://creativecommons.org/licenses/by-nc/3.0/) which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited.

Received 30 March 2021; Revised 6 May 2021; Accepted 14 June 2021 저자 직위: 김소라(석사 후 인턴연구원), 이수정(연구사), 양재형(연구사) https://doi.org/10.5657/KFAS.2021.0318

Korean J Fish Aquat Sci 54(3), 318-328, June 2021

동해안 참가자미의 해역별 산란특성 비교

319

Lee et al., 2006),

산란생태연구

(Chang et al., 2004; Cha et al., 2006)

가수행되어왔다

.

한편

,

국외연구로는연령과성장 연구

(Takahashi et al., 1995),

계절변화에따른섭이패턴연구

(Tominaga et al., 1995),

이석경에따른해역별성장차이연구

(Shimoda et al., 2006a),

^{성육장별치어의섭이및영양상태비} 교연구

(Kobayashi et al., 2015),

초기생활사단계에서의시공 간적인분포연구

(Joh and Wada, 2018)

등이있다

.

국내에서는 최근참가자미에관한연구가거의수행된바없으며

,

특히산 란생태연구는

15

년전연구결과이므로

,

현시점에서참가자미 의산란특성연구를통해과거선행연구와비교할필요가있을 것으로생각된다

.

따라서본연구에서는강원연안과경북연안에서식하고있 는참가자미의전장분포

,

성비

,

생식소숙도지수

,

난소의발달과 정

,

성숙체장등을분석하여최근참가자미의해역별산란생태 연구를수행하였고선행연구와의차이를살펴보았다

.

재료 및 방법

본연구는강원연안

(2018

년

1-12

월

)

과 경북연안

(2019

년

1-12

월

)

에서각각연안연승과연안자망에서매월어획된참가

자미를이용하였다

(Fig. 2).

^{채집된시료는실험실로옮겨암}

·

^수 성별을구분하였고

,

전장

(total length, TL)

은

0.1 cm

단위

,

체중

(body weight, BW)

과생식소중량

(gonad weight, GW)

은

0.01 g

까지측정하였다

.

본연구기간동안각해역의월간수온변화를알아보기위해 한국해양자료센터

(KODC)

^{의경북과강원연안의표층수온자} 료를사용하였다

(KODC, 2021).

참가자미의전장분포가암

·

수 간에차이가있는지알아보기위해

t-test

검정을하였고

,

월별

암

·

^{수성비의차이는}

chi-square test

^{를통해유의성검증을하} 였다

.

생식소숙도지수

(gonadosomatic index, GSI)

의월변화는해 Fig. 1. Annual variation of righteye flounders (Pleuronectidae) in the East sea from 1990 to 2019.

0 2,000 4,000 6,000 8,000 10,000 12,000 14,000

1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 2018 2019

1990s 2000s 2010s

Catch (M/T)

Year

25 20 15 10 5 0 5 10 15 20 25

14 16 18 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 Female N: 587 Mean: 23.2 cm

Male N: 434 Mean: 20.7 cm Gangwon

25 20 15 10 5 0 5 10 15 20 25

14 16 18 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 Female N: 952 Mean: 24.9 cm

Male N: 636 Mean: 22.0 cm Gyeongbuk

Frequency (%)

Total length (cm)

2018 2019

(A) (B) (C)

(D) (E) (F)

GVBD HY

PG CA Vtg1

Vtg2

Vtg3

POF

(A) (B) (C)

(D) (E) (F)

(G) (H) (I)

N

POF

NO CA

ZR FL YG

HY

GV

GVBD

0 10 20 30 40 50 60 70 80 90 100

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

Immature Maturing Mature Ripe/Spawning Spent 0 10 20 30 40 50 60 70 80 90 100

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

Immature Maturing Mature Ripe/Spawning Spent

0 10 20 30 40 50 60 70 80 90 100

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

Immature Maturing Mature Ripe/Spawning Spent

Gangwon Gyeongbuk

0 10 20 30 40 50 60 70 80 90 100

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

Immature Maturing Mature Ripe/Spawning Spent

Frequency (%)

2018 2019

Female

Male

0 25 50 75 100

0 5 10 15 20 25 30 35 40 45 50

0 25 50 75 100

0 5 10 15 20 25 30 35 40 45 50

Gangwon Gyeongbuk

Female

Proportion maturity (%)

Total length (cm) TL

_97.5%

= 29.7 cm

TL

_75%

= 25.8 cm

TL

_50%

= 24.2 cm

TL

_97.5%

= 25.7 cm

TL

_75%

= 19.3 cm

TL

_50%

= 16.5 cm 2018

0 25 50 75 100

0 5 10 15 20 25 30 35 40 45 50

TL

_97.5%

= 31.4 cm

TL

_75%

= 22.7 cm

TL

_50%

= 19.0 cm

2019

0 25 50 75 100

0 5 10 15 20 25 30 35 40 45 50

Male TL

_97.5%

= 26.4 cm

TL

_75%

= 21.8 cm

TL

_50%

= 19.8 cm

GSI

35 30 25 20 15 10 5 0 6 5 4 3 2 1 0

Gangwon Gyeongbuk

Female

Male

35 30 25 20 15 10 5 0 6 5 4 3 2 1 0

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec. Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

2018 2019

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec. Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

Fig. 2. Sampling area of the yellow striped flounder Pseudo- pleuronectes herzensteini in the coastal waters off Gangwon and Gyeongbuk.

0.0 5.0 10.0 15.0 20.0 25.0 30.0

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

Sea surface temperature (°C)

Gangwon (2018) Gyeongbuk (2019) Hatching water temperature (°C)

김소라

ㆍ

이수정

ㆍ

양재형

320

역별로구분하여다음의식으로구하였다

. GSI=

^GWBW^×100

여기서

GW

는생식소 중량

(g, wet weight), BW

는체중

(g, wet weight)

를나타낸다

.

참가자미의난모세포발달과정을관찰하기위해난소는측정 후절취하여

, Bouin’s solution

에

24

시간고정후흐르는담수에 수세하였다

.

^{이후난모세포관찰을용이하게하기위해조직표}

본은파라핀에포매하여

4-6 μm

두께로연속절편하였고

,

염색

은

hematoxylin-eosin (H-E)

을이용하였다

.

완성된조직표본 은광학현미경

(BX43; Olympus, Tokyo, Japan)

과이미지분석 소프트웨어

(TCapture Imaging Application; Tucsen, Fuzhou, China)

^{를사용하여관찰하였다}

.

난소발달과정은 육안관찰 및 조직학적 분석을병행하였고

,

정소발달과정은 육안관찰만을 참고하여 미숙

(immature),

중 숙

(maturing),

성숙

(mature),

완숙및방중

(ripe and spawning),

방후

(spent) 5

단계로구분하였다

(Murua et al., 2003; Brown- Peterson et al., 2011).

성숙전장은산란기로추정되는시기에중숙이상인개체를당 해년도에산란가능군으로가정하여산란가능한출현개체비 율을

logistic

식

(Zhang, 2010)

에적용하여추정하였다

.

P

_i

=

¹ 1+e

^(b

¹

^-b

²

^TL

ⁱ

⁾

여기서

P

_i는

i

전장계급에서의성숙비율

, TL

_i는

i

전장계급의 전장

, b

₁

, b

₂^{는상수이다}

.

또한

,

성숙전장은대상어종의자원상태를진단하기위한중요 한지표중하나이며

,

현재시행하고있는포획금지체장이적합 한지의여부를판단할수있는근거자료로사용될수있다

.

일

반적으로포획금지체장은

50%

^{성숙전장을기준으로하고있} 으나

,

본연구에서는참가자미의자원수준에적합한자원관리

방안을마련하기위해

50%, 75%, 97.5%

성숙전장을추정하여

과학적기준을제시하고자한다

.

결 과

전장분포

참가자미의전장분포는강원연안과경북연안에서어획된개 체를어업별로구분하여분석하였다

(Table 1, Fig. 3).

강원연안 에서어획된암컷참가자미는

587

^{마리였으며}

, 15.0-48.3 cm

^의 전장분포를보였고

,

평균전장은

23.2±6.0 cm

였다

.

또한수컷 참가자미는

434

마리가어획되었으며

, 15.0-36.0 cm

의전장분 포를보였고

,

평균전장은

20.7±3.9 cm

였다

.

경북연안에서어 획된암컷참가자미는

952

마리였으며

, 14.0-43.2 cm

의전장분 포를보였고

,

^{평균전장은}

24.9±4.0 cm

^였다

.

^{또한수컷참가자} 미는

636

마리가어획되었으며

, 16.2-31.2 cm

의전장분포를보 였고

,

평균전장은

22.0±2.3 cm

였다

.

강원연안에서채집된참 가자미의평균전장은암컷이수컷보다약

2.5 cm

컸으며

,

경북

연안또한암컷이수컷보다약

2.9 cm

크게나타나두해역모두

암

·

^{수전장간에차이가있는것으로나타났다}

(T-test, P<0.05).

암컷참가자미의평균전장은강원연안이경북연안보다약

1.7 cm

작았으며

,

수컷또한강원연안이경북연안보다약

1.3 cm

작게나타났다

.

성비

참가자미의성비는강원연안과경북연안을구분하여분석

하였다

(Table 1).

강원 연안에서 어획된 참가자미의 성비는

57.5:42.5

로암컷의비율이높았고

,

경북연안에서어획된참가

자미의성비또한

59.9:40.1

로암컷이수컷보다높은비율을보

여

,

두해역모두암컷의비율이수컷보다높은것으로나타났

Fig. 3. Length-frequency distribution of the yellow striped flounder Pseudopleuronectes herzensteini in the coastal waters off Gangwon and Gyeongbuk.

0 2,000 4,000 6,000 8,000 10,000 12,000 14,000

1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 2018 2019

1990s 2000s 2010s

Catch (M/T)

Year

25 20 15 10 5 0 5 10 15 20 25

14 16 18 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 Female N: 587 Mean: 23.2 cm

Male N: 434 Mean: 20.7 cm Gangwon

25 20 15 10 5 0 5 10 15 20 25

14 16 18 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 Female N: 952 Mean: 24.9 cm

Male N: 636 Mean: 22.0 cm Gyeongbuk

Frequency (%)

Total length (cm)

2018 2019

(A) (B) (C)

(D) (E) (F)

GVBD HY

PG CA Vtg1

Vtg2

Vtg3

POF

(A) (B) (C)

(D) (E) (F)

(G) (H) (I)

N

POF

NO CA

ZR FL YG

HY

GV

GVBD

0 10 20 30 40 50 60 70 80 90 100

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

Immature Maturing Mature Ripe/Spawning Spent 0 10 20 30 40 50 60 70 80 90 100

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

Immature Maturing Mature Ripe/Spawning Spent

0 10 20 30 40 50 60 70 80 90 100

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

Immature Maturing Mature Ripe/Spawning Spent

Gangwon Gyeongbuk

0 10 20 30 40 50 60 70 80 90 100

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

Immature Maturing Mature Ripe/Spawning Spent

Frequency (%)

2018 2019

Female

Male

50 75 100 0

25 50 75 100

0 5 10 15 20 25 30 35 40 45 50

Gangwon Gyeongbuk

Female

Proportion maturity (%)

TL

_97.5%

= 29.7 cm

TL

_75%

= 25.8 cm

TL

_50%

= 24.2 cm

TL

_97.5%

= 25.7 cm

TL

_75%

= 19.3 cm

TL

_50%

= 16.5 cm 2018

0 25 50 75 100

0 5 10 15 20 25 30 35 40 45 50

TL

_97.5%

= 31.4 cm

TL

_75%

= 22.7 cm

TL

_50%

= 19.0 cm

2019

50 75

100 TL

_97.5%

= 26.4 cm

TL

_75%

= 21.8 cm

TL

_50%

= 19.8 cm

GSI

35 30 25 20 15 10 5 0 6 5 4 3 2 1 0

Gangwon Gyeongbuk

Female

Male

35 30 25 20 15 10 5 0 6 5 4 3 2 1 0

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec. Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

2018 2019

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec. Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

동해안 참가자미의 해역별 산란특성 비교

321

다

(χ

²

-test, P<0.05).

생식소숙도지수의 월 변화

강원 연안에서 어획된암컷참가자미의 평균

GSI

는

1

월에

3.39±3.74

를보인후증가하여

2-5

월까지

4.66-7.96

을보이며 높은값을보였다

(Fig. 4).

이후

6

월부터감소하는경향을보이 나일부개체에서높은

GSI

^{가관찰되었다}

. 7-9

^월의평균

GSI

는

0.58-0.60

를보이며가장낮은값을보였다

. 10

월부터

GSI

가증가하면서

12

월까지평균

1.79-4.32

의

GSI

를나타냈다

.

강 원연안에서어획된수컷의평균

GSI

는

1-3

월에

1.05-1.16

을 보인후

4

월부터감소하는경향을보였지만

4-5

월에일부개체 에서높은

GSI

^{가관찰되었다}

. 7-10

^월의평균

GSI

^는

0.10-0.54

의낮은경향을보였으며

, 11

월과

12

월에는각각

1.72±1.12,

2.11±1.41

^{로증가하였다}

.

경북연안에서 어획된 암컷참가자미의 평균

GSI

는

1

월에

9.98±5.95

을보였고

2

월에

18.52±7.23

로가장높은값을나 타냈다

(Fig. 4).

이후평균

GSI

는

3

월부터

5

월까지상당히높 은값을유지하면서점차감소하는경향을보였다

. 6-9

월의평 균

GSI

^는

1.25-1.48

^{을보이며낮은경향을보였다}

. 10-12

^월에 는평균

GSI

값이

1.47-2.76

으로나타났으며

,

일부개체에서는

GSI

가증가하는경향을보였다

.

경북연안에서어획된수컷참

가자미의평균

GSI

는

1-2

월에

2.14-2.41

로가장높은값을나 타냈다

.

이후

6

월까지감소하기시작하였으며일부개체는평균 보다높은

GSI

^{값을보였다}

. 7-10

^월의평균

GSI

^는

0.11-0.27

^로 가장낮은경향을보였으며

, 11

월과

12

월에는각각

0.53±0.75, 1.56±2.22

으로증가하였다

.

Table 1. Monthly size distributions (total length, TL) and ratio of the yellow striped flounder Pseudopleuronectes herzensteini in the coastal waters off Gangwon and Gyeongbuk

Year Month Female Male

Range (cm) Mean n Sex ratio (%) Range (cm) Mean n Sex ratio (%)

Gangwon 2018

Jan. 16.0-33.6 22.0 51 50.0 16.5-26.5 18.6 51 50.0

Feb. 16.3-36.1 24.2 60 64.5 15.5-25.2 19.7 33 35.5

Mar. 15.1-37.4 20.2 38 43.2 15.1-31.6 21.2 50 56.8

Apr. 17.5-43.1 25.1 67 57.3 16.7-30.5 19.9 50 42.7

May 17.8-48.3 29.7 30 33.0 18.3-30.7 24.4 61 67.0

Jun. 15.3-36.3 24.3 63 79.7 15.3-29.5 21.2 16 20.3

Jul. 16.0-29.3 19.3 56 56.0 15.6-21.5 18.0 44 44.0

Aug. 15.0-23.8 18.0 30 68.2 15.2-30.6 19.3 14 31.8

Sep. 16.1-31.3 20.4 38 55.9 15.9-20.8 18.4 30 44.1

Oct. 15.7-42.5 25.6 34 49.3 15.0-31.0 21.3 35 50.7

Nov. 21.0-37.0 28.0 33 46.5 15.7-36.0 22.5 38 53.5

Dec. 17.8-38.8 22.6 87 87.9 18.3-28.0 22.2 12 12.1

All 15.0-48.3 23.2 587 57.5 15.0-36.0 20.7 434 42.5

Gyeongbuk 2019

Jan. 19.0-33.8 26.5 46 57.5 18.3-28.0 23.2 34 42.5

Feb. 15.5-42.3 26.0 102 63.8 16.2-28.5 20.4 58 36.3

Mar. 18.5-43.2 25.9 52 35.1 16.6-28.0 21.5 96 64.9

Apr. 14.0-35.5 25.1 71 75.5 17.0-25.8 22.0 23 24.5

May 20.8-36.1 25.8 115 75.7 21.3-31.2 24.6 37 24.3

Jun. 21.8-30.8 26.4 57 74.0 19.5-27.5 24.5 20 26.0

Jul. 18.8-30.7 23.1 77 48.4 17.8-27.0 21.8 82 51.6

Aug. 17.6-33.9 23.2 83 56.5 17.5-25.6 21.8 64 43.5

Sep. 19.5-33.8 23.5 45 43.7 18.3-26.5 22.0 58 56.3

Oct. 19.8-43.2 28.3 80 93.0 21.3-27.1 23.3 6 7.0

Nov. 17.7-28.3 22.0 107 55.4 17.0-29.4 21.1 86 44.6

Dec. 18.9-32.4 24.5 117 61.9 17.7-28.5 22.6 72 38.1

All 14.0-43.2 24.9 952 59.9 16.2-31.2 22.0 636 40.1

Total 14.0-48.3 24.1 1,539 59.0 15.0-36.0 21.5 1,070 41.0

김소라

ㆍ

이수정

ㆍ

양재형

322

Fig. 5. Photomicrographs of ovary stages of yellow striped flounder Pseudopleuronectes herzensteini. A, immature stage; B, maturing stage;

C, mature stage; D and E, ripe and spawning stage; F, spent stage; PG, primary growth stage oocyte; CA, cortical alveoli stage oocyte; Vtg1, primary vitellogenic oocyte; Vtg2, secondary vitellogenic oocyte; Vtg3, tertiary vitellogenic oocyte; GVBD, germinal vesicle breakdown;

HY, hydration; POF, postovulatory follicle complex. Scale bar=100 μm.

0 2,000 4,000 6,000 8,000 10,000 12,000 14,000

1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 2018 2019

1990s 2000s 2010s

Catch (M/T)

Year

25 20 15 10 5 0 5 10 15 20 25

14 16 18 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 Female N: 587 Mean: 23.2 cm

Male N: 434 Mean: 20.7 cm Gangwon

25 20 15 10 5 0 5 10 15 20 25

14 16 18 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 Female N: 952 Mean: 24.9 cm

Male N: 636 Mean: 22.0 cm Gyeongbuk

Frequency (%)

Total length (cm)

2018 2019

(A) (B) (C)

(D) (E) (F)

GVBD HY

PG CA Vtg1

Vtg2

Vtg3

POF

(A) (B) (C)

(D) (E) (F)

(G) (H) (I)

N

POF

NO CA

ZR FL YG

HY

GV

GVBD

0 10 20 30 40 50 60 70 80 90 100

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

Immature Maturing Mature Ripe/Spawning Spent 0 10 20 30 40 50 60 70 80 90 100

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

Immature Maturing Mature Ripe/Spawning Spent

0 10 20 30 40 50 60 70 80 90 100

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

Immature Maturing Mature Ripe/Spawning Spent

Gangwon Gyeongbuk

0 10 20 30 40 50 60 70 80 90 100

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

Immature Maturing Mature Ripe/Spawning Spent

Frequency (%)

2018 2019

Female

Male

0 25 50 75 100

0 5 10 15 20 25 30 35 40 45 50

0 25 50 75 100

0 5 10 15 20 25 30 35 40 45 50

Gangwon Gyeongbuk

Female

Proportion maturity (%)

Total length (cm) TL

_97.5%

= 29.7 cm

TL

_75%

= 25.8 cm

TL

_50%

= 24.2 cm

TL

_97.5%

= 25.7 cm

TL

_75%

= 19.3 cm

TL

_50%

= 16.5 cm 2018

0 25 50 75 100

0 5 10 15 20 25 30 35 40 45 50

TL

_97.5%

= 31.4 cm

TL

_75%

= 22.7 cm

TL

_50%

= 19.0 cm

2019

0 25 50 75 100

0 5 10 15 20 25 30 35 40 45 50

Male TL

_97.5%

= 26.4 cm

TL

_75%

= 21.8 cm

TL

_50%

= 19.8 cm

GSI

35 30 25 20 15 10 5 0 6 5 4 3 2 1 0

Gangwon Gyeongbuk

Female

Male

35 30 25 20 15 10 5 0 6 5 4 3 2 1 0

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec. Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

2018 2019

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec. Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

Fig. 4. Monthly variation of GSI (gonadosomatic index) of the yellow striped flounder Pseudopleuronectes herzensteini in the coastal waters off Gangwon and Gyeongbuk.

0 2,000 4,000 6,000 8,000 10,000 12,000 14,000

1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 2018 2019

1990s 2000s 2010s

Catch (M/T)

Year

25 20 15 10 5 0 5 10 15 20 25

14 16 18 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 Female N: 587 Mean: 23.2 cm

Male N: 434 Mean: 20.7 cm Gangwon

25 20 15 10 5 0 5 10 15 20 25

14 16 18 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 Female N: 952 Mean: 24.9 cm

Male N: 636 Mean: 22.0 cm Gyeongbuk

Frequency (%)

Total length (cm)

2018 2019

(A) (B) (C)

(D) (E) (F)

GVBD HY

PG CA Vtg1

Vtg2

Vtg3

POF

(A) (B) (C)

(D) (E) (F)

(G) (H) (I)

N

POF

NO CA

ZR FL YG

HY

GV

GVBD

0 10 20 30 40 50 60 70 80 90 100

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

Immature Maturing Mature Ripe/Spawning Spent 0 10 20 30 40 50 60 70 80 90 100

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

Immature Maturing Mature Ripe/Spawning Spent

0 10 20 30 40 50 60 70 80 90 100

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

Immature Maturing Mature Ripe/Spawning Spent

Gangwon Gyeongbuk

0 10 20 30 40 50 60 70 80 90 100

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

Immature Maturing Mature Ripe/Spawning Spent

Frequency (%)

2018 2019

Female

Male

0 25 50 75 100

0 5 10 15 20 25 30 35 40 45 50

0 25 50 75 100

0 5 10 15 20 25 30 35 40 45 50

Gangwon Gyeongbuk

Female

Proportion maturity (%)

Total length (cm) TL

_97.5%

= 29.7 cm

TL

_75%

= 25.8 cm

TL

_50%

= 24.2 cm

TL

_97.5%

= 25.7 cm

TL

_75%

= 19.3 cm

TL

_50%

= 16.5 cm 2018

0 25 50 75 100

0 5 10 15 20 25 30 35 40 45 50

TL

_97.5%

= 31.4 cm

TL

_75%

= 22.7 cm

TL

_50%

= 19.0 cm

2019

0 25 50 75 100

0 5 10 15 20 25 30 35 40 45 50

Male TL

_97.5%

= 26.4 cm

TL

_75%

= 21.8 cm

TL

_50%

= 19.8 cm

GSI

35 30 25 20 15 10 5 0 6 5 4 3 2 1 0

Gangwon Gyeongbuk

Female

Male

35 30 25 20 15 10 5 0 6 5 4 3 2 1 0

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec. Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

2018 2019

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec. Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

난소의 발달단계

미숙 단계의 난소

(Fig. 5A)

^{에는 초기성장기 난모세포}

(63.8

±16.8 μm)

가관찰되었다

(Fig. 6A).

초기성장기난모세포는핵

의주변부에다수의인이관찰되었으며

,

^{모든발달단계의난소} 에서연중관찰되는모습을보였다

.

중숙단계의난소

(Fig. 5B)

에는초기성장기난모세포가성장함에따라난모세포의크기가

동해안 참가자미의 해역별 산란특성 비교

323

132.7±14.3 μm

로커졌고세포질의바깥부분에붉게염색된

난황포들이관찰되었다

(Fig. 6B).

성숙단계의난소

(Fig. 5C)

에 는난황형성기에접어들며세포질내에호산성의난황구가축 적되었으며핵의주변부쪽으로넓게형성되는경향을보였다

(153.2±21.6 μm) (Fig. 6C).

^{또한난모세포가발달함에따라} 난황구가핵주변을제외하고균일하게분포하였고난모세포 의크기가

224.1±29.4 μm

로커졌다

(Fig. 6D).

완숙및방중단 계의난소

(Fig. 5D, 5E)

에는난모세포가

445.0±39.9 μm

로발 달하며핵주위까지난황구가고르게분포하였고

,

잘발달된방 사대가뚜렷하게관찰되었다

(Fig. 6E).

^{난모세포가최종성숙} 기에접어들면서핵은동물극쪽으로이동하는모습을보였으 며

(502.8±35.4 μm) (Fig. 6F),

곧이어핵막이불분명해지며세 포질과염색질이 섞이는난핵포붕괴

(germinal vesicle break- down, GVBD)

가관찰되었다

(551.4±63.6 μm) (Fig. 6G).

이 후

,

^{배란직전의수화된난모세포가보였다}

(585.4±66.1 μm) (Fig. 6H).

방후단계

(Fig. 5F)

의난소에는배란후수화된난모 세포를둘러싸고있던여포세포층즉

,

배란후여포세포

(post- ovulatory follicle complex, POF)

와초기성장기난모세포가관 찰되었다

(Fig. 6I).

성숙도의 월 변화

난소발달과정은육안관찰과조직학적분석을병행하였고

,

^정 소발달과정은육안관찰만을참고하여성숙도의월변화를알

아보았다

(Fig. 7).

강원연안에서어획된암컷은미숙단계가연

중관찰되었으며

,

특히

, 7-9

월에는

100%

로나타났다

.

중숙단 계는

1-5

월까지나타났으며

, 7-9

월에관찰되지않다가

10

월부 터출현하였다

.

^{성숙단계는주로}

2-6

^{월에나타났으며}

,

^완숙및 방중단계는

4-5

월에

10.0-10.4%

로출현하였다

.

방후단계는

4-6

월에

25.4-36.7%

로출현하였다

.

강원연안에서어획된수컷 또한미숙단계가연중관찰되었으며

, 7-10

월에

97.1-100%

로 가장높은비율을보였다

.

중숙단계는

1-4

월에걸쳐나타났으 며

, 7-9

^{월에관찰되지않다가}

10

^{월부터출현하기시작하였다}

.

성숙단계는

2-5

월에나타났으며

,

완숙및방중단계는

4-5

월 에

2.0-18.0%

로출현하였다

.

방후단계는

4-6

월에

10.0-43.8%

로관찰되었다

.

경북연안에서어획된암컷은미숙단계가

1-6

월에

2.2-29.3%

로낮게나타났으며

,

^이후

7-12

^월까지

56.3-97.4%

^{로출현하였} 다

.

중숙단계는

1-2

월에나타났으며

, 2-4

월에는관찰되지않 Fig. 6. Photomicrographs of oocyte development stages of yellow striped flounder Pseudopleuronectes herzensteini. A, primary growth stage oocyte (PG); B, cortical alveoli stage oocyte (CA); C, primary vitellogenic oocyte (Vtg1); D, secondary vitellogenic oocyte (Vtg2);

E, tertiary vitellogenic oocyte (Vtg3); F, germinal vesicle migration (GVM); G, germinal vesicle breakdown (GVBD); H, hydration (HY);

I, postovulatory follicle complex (POF); N, nucleus; NO, nucleolus; YG, yolk globule; ZR, zona radiate; FL, follicle layer; GV, germinal vesicle. Scale bar=100 μm.

0 2,000 4,000 6,000 8,000 10,000 12,000 14,000

1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 2018 2019

1990s 2000s 2010s

Catch (M/T)

Year

25 20 15 10 5 0 5 10 15 20 25

14 16 18 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 Female N: 587 Mean: 23.2 cm

Male N: 434 Mean: 20.7 cm Gangwon

25 20 15 10 5 0 5 10 15 20 25

14 16 18 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 Female N: 952 Mean: 24.9 cm

Male N: 636 Mean: 22.0 cm Gyeongbuk

Frequency (%)

Total length (cm)

2018 2019

(A) (B) (C)

(D) (E) (F)

GVBD HY

PG CA Vtg1

Vtg2

Vtg3

POF

(A) (B) (C)

(D) (E) (F)

(G) (H) (I)

N

POF

NO CA

ZR FL YG

HY

GV

GVBD

0 10 20 30 40 50 60 70 80 90 100

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

Immature Maturing Mature Ripe/Spawning Spent 0 10 20 30 40 50 60 70 80 90 100

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

Immature Maturing Mature Ripe/Spawning Spent

0 10 20 30 40 50 60 70 80 90 100

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

Immature Maturing Mature Ripe/Spawning Spent

Gangwon Gyeongbuk

0 10 20 30 40 50 60 70 80 90 100

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

Immature Maturing Mature Ripe/Spawning Spent

Frequency (%)

2018 2019

Female

Male

0 25 50 75 100

0 5 10 15 20 25 30 35 40 45 50

0 25 50 75 100

0 5 10 15 20 25 30 35 40 45 50

Gangwon Gyeongbuk

Female

Proportion maturity (%)

Total length (cm) TL

_97.5%

= 29.7 cm

TL

_75%

= 25.8 cm

TL

_50%

= 24.2 cm

TL

_97.5%

= 25.7 cm

TL

_75%

= 19.3 cm

TL

_50%

= 16.5 cm 2018

0 25 50 75 100

0 5 10 15 20 25 30 35 40 45 50

TL

_97.5%

= 31.4 cm

TL

_75%

= 22.7 cm

TL

_50%

= 19.0 cm

2019

0 25 50 75 100

0 5 10 15 20 25 30 35 40 45 50

Male TL

_97.5%

= 26.4 cm

TL

_75%

= 21.8 cm

TL

_50%

= 19.8 cm

GSI

35 30 25 20 15 10 5 0 6 5 4 3 2 1 0

Gangwon Gyeongbuk

Female

Male

35 30 25 20 15 10 5 0 6 5 4 3 2 1 0

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec. Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

2018 2019

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec. Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

김소라

ㆍ

이수정

ㆍ

양재형

324

다가

5

월부터다시출현하기시작하였다

.

성숙단계는

1-5

월까 지나타났고

,

완숙및방중단계는

1-4

월까지출현하였으며주 로

2-3

^월에

20.3-43.5%

^{로관찰되었다}

.

^{방후단계는}

1-8

^월까지 나타났으며

,

주로

4-5

월에

57.7-73.7%

로높은비율로관찰되었

다

.

경북연안에서어획된수컷은미숙단계가

3-12

월까지나타

났으며

,

특히

7-11

월에는

94.3-100%

로가장높은비율을보였 다

.

중숙단계는

5-6

월

, 11-1

월에나타났으며

, 6

월과

12

월에는 각각

20.0%, 52.5%

^{의비율을보였다}

.

^{성숙단계는}

1-6

^월까지 출현하였고

,

완숙및방중단계는

1

월과

3-4

월에나타났으며특 히

1

월과

3

월에는각각

38.2%

와

48.6%

로관찰되었다

.

방후단 계는

3-6

월까지나타났다

.

성숙전장

강원연안과경북연안에서어획된암

·

^{수참가자미의}

50%, 75%, 97.5%

성숙전장을알아보았다

(Fig. 8).

강원연안에서어 획된암컷참가자미의

50%, 75%, 97.5%

성숙전장은순차적으 로

24.2 cm, 25.8 cm, 29.7 cm

였고

,

수컷참가자미는

19.8 cm, 21.8 cm, 26.4 cm

로나타났다

.

반면

,

경북연안에서어획된암 컷참가자미의

50%, 75%, 97.5%

^{성숙전장은순차적으로}

19.0 cm, 22.7 cm, 31.4 cm

였고

,

수컷참가자미는

16.5 cm, 19.3 cm,

25.7 cm

로나타났다

.

강원연안과경북연안모두암컷의

50%

성숙전장이수컷보다각각

4.4, 2.5 cm

컸으며

,

강원연안에서 어획된참가자미는경북연안에서어획된참가자미보다암컷 과수컷에서각각

5.2, 3.3 cm

큰것으로나타났다

.

따라서본연 구에서는참가자미암컷이수컷보다성숙전장이크며

,

강원에 서어획된참가자미가경북에서어획된참가자미보다성숙전장 이큰것으로나타났다

.

고 찰

동해안에서식하는가자미과어류에서는암컷이수컷보다성 장이빠르고

(Choe et al., 1999; Lee et al., 2006; Yang et al., 2017),

크기도더큰것으로보고되고있다

(Cha et al., 2008;

Choi et al., 2013; Kim et al., 2016; Yang et al., 2018).

^본연구 에서또한강원과경북연안에서식하는참가자미의암

·

수간에 전장차이를보였다

.

강원연안에서채집된참가자미는경북연안에서채집된참 가자미보다상대적으로작은전장이주로어획되었다

.

이처 럼조사해역별로전장에차이가나타나는이유는서식지및조 사시기에따른개체군의밀도

,

수온

,

먹이생물의풍도등의차

0 2,000 4,000 6,000 8,000 10,000 12,000 14,000

1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 2018 2019

1990s 2000s 2010s

Catch (M/T)

Year

25 20 15 10 5 0 5 10 15 20 25

14 16 18 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 Female N: 587 Mean: 23.2 cm

Male N: 434 Mean: 20.7 cm Gangwon

25 20 15 10 5 0 5 10 15 20 25

14 16 18 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 Female N: 952 Mean: 24.9 cm

Male N: 636 Mean: 22.0 cm Gyeongbuk

Frequency (%)

Total length (cm)

2018 2019

(A) (B) (C)

(D) (E) (F)

GVBD HY

PG CA Vtg1

Vtg2

Vtg3

POF

(A) (B) (C)

(D) (E) (F)

(G) (H) (I)

N

POF

NO CA

ZR FL YG

HY

GV

GVBD

0 10 20 30 40 50 60 70 80 90 100

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

Immature Maturing Mature Ripe/Spawning Spent 0 10 20 30 40 50 60 70 80 90 100

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

Immature Maturing Mature Ripe/Spawning Spent

0 10 20 30 40 50 60 70 80 90 100

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

Immature Maturing Mature Ripe/Spawning Spent

Gangwon Gyeongbuk

0 10 20 30 40 50 60 70 80 90 100

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

Immature Maturing Mature Ripe/Spawning Spent

Frequency (%)

2018 2019

Female

Male

0 25 50 75 100

0 5 10 15 20 25 30 35 40 45 50

0 25 50 75 100

0 5 10 15 20 25 30 35 40 45 50

Gangwon Gyeongbuk

Female

Proportion maturity (%)

Total length (cm) TL

_97.5%

= 29.7 cm

TL

_75%

= 25.8 cm

TL

_50%

= 24.2 cm

TL

_97.5%

= 25.7 cm

TL

_75%

= 19.3 cm

TL

_50%

= 16.5 cm 2018

0 25 50 75 100

0 5 10 15 20 25 30 35 40 45 50

TL

_97.5%

= 31.4 cm

TL

_75%

= 22.7 cm

TL

_50%

= 19.0 cm

2019

0 25 50 75 100

0 5 10 15 20 25 30 35 40 45 50

Male TL

_97.5%

= 26.4 cm

TL

_75%

= 21.8 cm

TL

_50%

= 19.8 cm

GSI

35 30 25 20 15 10 5 0 6 5 4 3 2 1 0

Gangwon Gyeongbuk

Female

Male

35 30 25 20 15 10 5 0 6 5 4 3 2 1 0

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec. Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

2018 2019

Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec. Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.

Fig. 7. Monthly changes in maturity stages female and male of the yellow striped flounder Pseudopleuronectes herzensteini in the coastal waters off Gangwon and Gyeongbuk.