Energy Budget of Spiny Lebbeid Shrimp Lebbeus groenlandicus Larvae

(1)

801

서 론

동물의에너지수지에관한연구는에너지흐름을이해하고동 물생태계의기능을파악하는데있어유용한수단이된다

.

^생태 계는복잡한먹이사슬또는먹이그물과같은영양적구조로집 단간의에너지흐름의양에밀접하게관련되어있으며

,

^이러한 복잡한먹이망속에서그물질의전환에따른에너지흐름을 파악하기위해서에너지수지를분석하는것이매우중요하다

(Klekowski and Duncan, 1975).

^어느^한^생물의^개체에^유입되 는모든에너지는열로써상실하거나

,

^다른^영양^단계로^전이된 다

.

^생물학적^물질계는^{물질전환에}^따른^에너지^흐름에^의해서 체계화되어있다

.

그리고먹이사슬로또는먹이그물과같은영 양적구조는집단의에너지흐름의양에밀접하게관련된다

.

^그 러므로에너지흐름의파악은자연집단의에너지수지를분석하 는것이필요하지만

,

^우선^개체의^생활사에^따른^{에너지수지를} 정확하게분석하는것이매우중요하다

(Park, 1989).

갑각류의에너지수지에관해서는자연과실험실내집단을대 상으로많은연구가진행되어있으며

(Clutter and Theilacker, 1971; Levine and Sulkin, 1979; Emmerson, 1984; Harms, 1987; Kim and Chin, 1991),

^성장

,

^산소소비

,

^질소배설^및^섭취 에대한에너지함량을파악하여에너지수지방식을통해서그 영양적체계를구성해볼수있다

(Park, 1989).

^동물의^에너지 수지에관한연구는다른영양단계간의에너지전이유형을이 해하는총체적지표가되며

,

^그^동물의^생산성과^생태적^효율을 예측하는자료로서유용하다

(Vernberg and Vernverg, 1981).

가시배새우

, Lebbeus groenlandicus

는십각목

(Order Decap- oda)

^{꼬마새우과}

(Family Hippolytidae)

^에^속하는^종으로^꼬마 새우과중에서도거대종이며

,

특히한국산꼬마새우과중에서 는가장크다

.

^몸이^짧고^굵으며

,

^두흉갑이^짧고^거친^털로^덮여 있다

.

^이^종은^울릉도^근해에서^통발로^주로^{어획하는데}

,

^수심

200-400 m

^에^주로^서식한다

.

^한해성^종으로서^아한대^해역에

분포하고있다

.

^우리나라^동해

,

^그린란드

,

^미국^동북부

,

^캐나다

Article history;

Received 8 May 2013; Revised 27 August 2013; Accepted 29 October 2013

*Corresponding author: Tel: +82. 33. 640. 2342 Fax: +82. 33. 640. 2340 E-mail address: [email protected]

Kor J Fish Aquat Sci 46(6) 801-806, December 2013 http://dx.doi.org/10.5657/KFAS.2013.0801 pISSN:0374-8111, eISSN:2287-8815

ⓒ The Korean Society of Fishereis and Aquatic Science. All rights reserved

가시배새우(Lebbeus groenlandicus)의 유생 단계 에너지수지

김원기·권오남

¹

·박기영

^2*

양양군청 해양수산과, ¹강릉원주대학교 해양생물연구교육센터, ^*2강릉원주대학교 해양자원육성학과

Energy Budget of Spiny Lebbeid Shrimp Lebbeus groenlandicus Larvae

Won Gi Kim, O-Nam Kwon¹ and Kie-Young Park^2*

Marine and Fisheries Division, Yangyang country Office, Yangyang-gun 215-701, Korea

1

Marine Biology Center for Research and Education, Gangneung-Wonju National University, Gangneung 210-853, Korea

2

Department of Marine Bioscience, Gangneung-Wonju National University, Gangneung 210-702, Korea

Spiny lebbeid shrimp Lebbeus groenlandicus larvae were reared in the laboratory to estimate the energy budget from the zoeal to the post-larval stage. Energy expended by larvae on growth and respiration was determined from values for feeding, growth, molting, and metabolism. We calculated that 16.22 J were used for growth throughout all larval stages. Energy loss during molt- ing was estimated as 1.03 J, and energy used for respiration was estimated as 1.31 J. Energy taken in by feeding was estimated as 77.16 J, while the sum of energies expended in excretion and egestion was 58.61 J. Larvae were estimated to assimilate 24.6%

of ingested food as energy and to use ~85% of the assimilated energy for somatic growth. Gross growth efficiency (K

₁

) and net growth efficiency (K

₂

) were shown to be ~22% and 93%, respectively. Maintenance costs of respiration were estimated as ~9.7%

of assimilated energy.

Key words: Lebbeus groenlandicus, Metabolism, Energy budget, Spiny lebbeid shrimp, Assimilation efficiency, Growth efficiency

(2)

북극해연해

,

^{오호츠크해}

,

^일본^홋카이도^등^{전세계적으로}^널리 분포하고있다

(NFRDI, 2007).

따라서본연구는가시배새우유생을대상으로성장

,

탈피

,

호 흡

,

^섭취^및^에너지^함량을^측정하여^유생^발달^동안^에너지수 지를분석하고섭취와성장에따른유생의에너지흐름의경향 을파악함으로써개체및개체군에대한생리적기초자료를얻 는데그목적이있다

.

재료 및 방법

실험동물 및 유생사육

실험에 사용한 가시배새우 유생은 평균 체장

117.22 mm,

중량

38.65 g

^의^{모하로부터}^얻었다

.

^내장을^제거한^민들조개

, Macridiscus melanaegis

를부화직전인암적색의알을달고있 는모하들에게먹이로공급하였다

.

^이들^모하들을^별도^용기에 수용하여부화당일혹은부화후

1

^일령의^부유성

zoea I

^기^유생 을수집하여

8±0.3℃

^에서^실험에^{사용하였다}

.

성장 및 에너지함량

초기유생의개체별성장을측정하기위하여수온

8±0.3℃,

250 mL

^용기

(

^사육수

, 200 mL)

^에^수용한^후^부화^직후^유생을

1

^마리씩^수용하여

Artemia nauplii (

^이하

Artemia)

^를^약

10

^개 체

/mL

^의^농도로^{공급하면서}^매일^아침^탈피체를^수거하여^크 기를측정하였다

.

성장측정은유생의손상을방지하기 위해 실험중인유생의탈피체를수거하여 측정하고

,

^각^유생단계 의탈피체의에너지함량및건조중량은각 발달단계별로대

형사육

(1 ton)

수조에서무작위추출하여측정하였다

.

실험유

생의체장은마지막복부체절의말단에서두흉갑의 앞쪽끝 까지의직선거리를광학현미경

(OLYMPUS IX70)

^을^이용하여

0.01 mm

단위로측정하였다

.

또한초기유생단계별유생및탈 피체에에너지함량은자동폭발열량계

(Van calorimeter model PARR1315, USA)

^로^측정하여

‘J'

^로^{표시하였다}

.

산소소비

유생의산소소비량은

250 mL

^산소병에^{가시배새우}^부화^직후 유생을

5

^마리씩^수용한^후

,

^실험^전후^각^{용존산소를}

DO meter (YSI 556MPS, USA)

로측정하였고

,

실험전후의용존산소차 로써정량하였다

.

^각^실험은

5

^회^반복한^평균치로^개체당^단위 시간당산소소비량을

μLO

₂

/ind/day

^로^{표시하였다}

.

^{산소소비량} 은대사열생산에의해소비된에너지로전환시키기위하여

Gn-

aiger (1983)

^가^이용한^{산소분자의}^에너지^{전화계수를}^이용하

였다

(1 mgO

₂

= 3.38 cal).

^대사에^쓰인^에너지는

4.71×10

^-3

cal/

μLO

₂의변환계수를사용하여계산하였다

.

개체의산소소비량 은환경수의산소분압에영향을받기때문에

(Ikeda, 1974),

^산 소소비측정이끝날때의실험용기내용존산소량이실험시작

시산소량의

60%

^이하로^떨어지지^않게^{조절하였다}

.

먹이 섭취

유생의섭취량측정시에는

250 mL

^의^용기에^{가시배새우}^유

생을

1

^마리씩^수용한^후^일정량의^계수된

Artemia nauplius

^를 투여하여

24

^시간^동안의^섭취량을^{측정하였다}

.

^그리고

1

^개체당 일간

Artemia nauplii

섭취량은

no./ind./day

으로나타내었다

.

섭취에너지는

1

^마리가

1

^일^섭취한

Artemia nauplii

의개체당 중량

(Dry weight 1.97 μg/nauplius, 4,619 cal/mg) (Emmerson, 1980)

^과

Artemia nauplii 1

^마리의^{에너지함량}

(37.9 mJ) (Park, 1989)

을기준으로계산하였다

.

에너지 수지

각유생단계및전체유생발생에대한에너지수지는

Standard IBP

^방정식

(Klekowski and Duncan, 1975)

^으로^{나타내었다}

.

C = Pg + Pe + R + E ………①

①

식에

C

^는^{섭취에너지}

, Pg

^는^{유생단계별}^체성장에^쓰인^에 너지

, Pe

는탈피에쓰인에너지

, R

은호흡에쓰인대사에너지

, E

^는^{대사산물인}^뇨^{배설에너지}^그리고^대사되지^않고^배출되는 분배출에너지

,

섭취에너지

(C)

와동화에너지

(A = Pg + Pe + R)

간의차로서나타내었다

(Dawirs, 1983).

생태효율

수산동물의동화효율측정은먹이섭취량과분배출량을측정 하여먹이섭취량에대한동화량을백분율로하여동화효율을 구하였다

.

^{가시배새우의}^유생에^대한^섭취

,

^성장

,

^탈피^및^호흡 을측정한자료를사용하여다음효율들을계산하였다

.

동화효율

: (Pg + Pe + R) / C × 100 ………②

②

식은섭취량중동화된양의백분율을나타낸다

.

총성장효율

(K1): (Pg + Pe) / C × 100 ………③

③

식은섭취량중성장으로전환된양의백분율을나타낸다

.

순성장효율

(K2): (Pg + Pe) / (Pg + Pe + R) × 100 ………④

④

식은동화량중성장으로전환된양의백분율을나타낸다

.

결 과

성장 및 에너지 함량

가시배새우의 부화후 경과시간에 따른 유생

(zoea I, II

^및

megalopa)

의체장

,

건조중량및유생과탈피체의에너지함량

은

Table 1

^에^{나타내었다}

. Zoea I

^기에서

megalopa

^기까지의^체 장성장은지수적인증가를보였다

(Fig. 1).

성장에사용된에너지즉

,

^생체^에너지^함량은

J/ind.

^로^산출 하였다

. ZoeaⅠ

^기에서

megalopa

^기까지^성장하는^동안

mega-

lopa

기의생체에너지는

16.21 J

의에너지를체성장에사용하 였다

.

^성장에^수반되어^탈피체로 ^손실된^에너지는^탈피체^에

(3)

너지함량을

J/molting body

^로 ^{계산하였는데}

, zoeaⅠ

^기부터

megalopa

기까지유생

1

마리가사용한탈피에너지는

1.03 J

이

었다

(Table 1).

^{체장성장과}^{건조중량은}^경과^일수에^대해서^각

각

y=6.088e

^0.116x

(R

²

=0.973)

^와

y=0.165e

^0.468x

(R

²

=0.893)

^로^나 타났다

(Fig. 1

과

2).

산소소비

가시배새우유생발달단계에따라산소소비량은

zoeaⅠ

^기의

1.449 μLO

₂

/ind./day

^에서

megalopa

^기의

4.731 μLO

₂

/ind,/day

로성장에따라 증가하였다

(Table 2).

^{일간대사에너지}

(J/ind./

day)

^를 ^사용하여 ^{유생단계별}^{경과시간에} ^따른 ^{대사에너지를} 계산하면

zoeaⅠ

기기간동안에는

0.268 J, zoeaⅡ

기는

0.601 J, megalopa

^기^기간^동안에는

0.982 J

^을^소비하여^유생기간^동 안소비된총산소소비량은

1.851 J

^이었다

.

섭취

가시배새우의

zoea

유생들과

megalopa

유생들에 의해 섭 취된일간섭취량과일간섭취에너지를

Table 3

. ZoeaⅠ

^기의

29.6 Artemia/ind./day

^에서

megalopa

^기의

127.7 Artemia/ind./day

까지유생발달의진행과함께먹이섭취량은

급격히증가하였다

. ZoeaⅠ

^기는

1.12 J/ind./day

^의^에너지를^섭 취하는것으로조사되었으며

,

유생이성장함에따라섭취에너 지는증가하여

megalopa

^기는

4.84 J

^의^에너지를^{섭취하였다}

.

각유생단계별경과시간에따른각단계의섭취에너지를바탕 으로전유생기동안섭취한총섭취에너지는

80.09 J

이었다

. 에너지수지

성장에따른전유생기동안의유생의성장에너지함량

(Pg)

^및 탈피체의에너지함량

(Pe)

^은

Table 1

^에서^제시한^결과를^사용하 였으며

,

^호흡^{대사에너지}

(R)

^는

Table 2

^의^{자료로부터}^계산하였 다

.

^그리고^{섭취에너지}

(C)

^는

Table 3

^의^일간^섭취량과^일간^섭 취에너지로부터계산하였다

.

뇨배설과분배출에너지

(E)

는섭 취에너지

(C)

^와^{동화에너지}

(A)

^의^차이로서^{계산하였다}

.

^전^유 생기동안의누적에너지수지를

Table 4

.

^가시배 새우유생의섭취

,

^탈피체

,

^호흡^및^배설^에너지는^성장과^함께 Fig. 1. Relationship between intermolt period and body length of

Lebbeus groenlandicus for the different larval stages.

Fig. 2. Relationship between intermolt period and dry weight of Lebbeus groenlandicus for the different larval stages.

Table 1. The intermolt periods, body length, body weight and energy contents of Lebbeus groenlandicus for the different larval stages

Larval

stage Intermolt periods (day)

Body length

(mm)

weight Dry

(mg) J/ind. J/molt

ZoeaⅠ 0 6.92 0.29 16.21 1.03

ZoeaⅡ 8.32 7.52 0.35 16.21 1.03

Megalopa 10.98 8.74 0.74 16.21 1.03

Table 2. Individual oxygen consumption and respiratory energy expenditure of Lebbeus groenlandicus for the different larval stages

Larval stage O2 consumption Respiratory energy expenditure μLO²/ind./day J/ind./day J/whole

stages

Zoea Ⅰ 1.449 0.032 0.268

Zoea Ⅱ 2.467 0.055 0.601

Megalopa 4.731 0.105 0.982

ZoeaⅠ-

Megalopa 8.647 0.192 1.851

1 mgO₂= 3.38 cal (Gnaiger, 1983) 1 cal = 4.71×10^-3 J

y = 6.088e

^0.116x

R

²

= 0.973 10

9 8 7 6

5 0 8.32 19.30

Body length (mm)

Intermolt periods (day)

Zoea I

Zoea II

Megalopa

y = 0.165e

^0.468x

R

²

= 0.893 1.0

0.9 0.7 0.5 0.3 0.1 0.8 0.6 0.4 0.2

0.0 0 8.32 19.30

Body length (mm)

Intermolt periods (day)

Zoea I

Zoea II

Megalopa y = 6.088e

^0.116x

R

²

= 0.973 10

9 8 7 6

5 0 8.32 19.30

Body length (mm)

Intermolt periods (day)

Zoea I

Zoea II

Megalopa

y = 0.165e

^0.468x

R

²

= 0.893 1.0

0.9 0.7 0.5 0.3 0.1 0.8 0.6 0.4 0.2

0.0 0 8.32 19.30

Body length (mm)

Intermolt periods (day)

Zoea I

Zoea II

Megalopa

(4)

증가하였다

. ZoeaⅠ

기에서

megalopa

기까지의총누적섭취에 너지는

80.09 J

^이었으며

,

^이것은^성장에

16.21 J,

^탈피체^손실 에

1.03 J

^그리고^호흡에

1.85 J

^의^에너지를^사용하며^뇨와^분

으로

61.00 J

의에너지가배설과배출이이루어지는것으로조

사되었다

. 생태효율

가시배새우유생의섭취

,

^성장

,

^탈피^및^호흡을^측정한^자료 로부터생태효율을

Table 5

.

^유생의^{동화효율은}

19.09%

이었으며

,

총성장효율

(K

₁

)

은

21.53%,

순성장효율

(K

₂

)

은

90.30%

^로^나타났다

.

^각^{에너지수지}^{구성요소들의}^섭취에너

지에대한백분율은성장이

20.24%,

^탈피체가

1.29%,

^대사에 너지가

2.31%

그리고뇨와분으로

76.16%

를각각차지하였다

.

ZoeaⅠ

^기부터

megalopa

^기까지의^총^{동화에너지는}^전체^섭취

에너지의

23.84%

인

19.09 J

이었으며

,

각에너지수지구성요 소들이동화에너지에대한백분율은성장이

84.91%,

^탈피체가

5.40%

^그리고^{대사에너지가}

9.70%

^를^각각^{차지하였다}

.

^또한

zoeaⅠ

기에서

megalopa

기까지의 총생산에너지는총동화에

Table 3. Consumption rate and consumption energy of Lebbeus groenlandicus for the different larval stages

Larval stage Feed consumption rate Feed consumption energy

Artemia nauplii

ind./ind./day Artemia nauplii

μg/ind./day J/ind./day J/ whole stages

Zoea Ⅰ 29.6 58.31 1.12 9.33

Zoea Ⅱ 61.3 120.75 2.32 25.51

Megalopa 127.7 251.56 4.84 45.25

ZoeaⅠ-Megalopa 218.6 430.62 8.28 80.09

37.9 mJ/Artemia nauplius (Park, 1989)

Table 4. Energy budgets of Lebbeus groenlandicus for the different larval stages

Larval stage C Pg Pe R E

Energy budget per hour (mJ)

Zoea Ⅰ 46.72 9.77 0.60 1.34 35.02

Zoea Ⅱ 96.80 18.52 1.18 2.28 74.83

Megalopa 201.65 41.80 2.67 4.38 152.80

Energy budget for the each larval and total development of zoeaⅠ to megalopa (J)

Zoea Ⅰ 9.33 1.95 0.12 0.27 6.99

Zoea Ⅱ 25.51 4.88 0.31 0.60 19.72

Megalopa 45.25 9.38 0.60 0.98 34.29

ZoeaⅠ~ Megalopa 80.09 16.21 1.03 1.85 61.00

Table 5. Accumulative energy budget of Lebbeus groenlandicus larvae

Symbol Total energy (J) %P %A %C

Molting Pe 1.03 5.97 5.40 1.29

Growth Pg 16.21 94.03 84.91 20.24

Total production P 17.24 100.00 90.30 21.53

Respiration R 1.31 9.70 2.31

Total assimilation A 19.09 100.00 23.84

Excretion & Egestion E 61.00 76.16

Consumption C 80.09 100.00

%P : Total production percentage, %A : Total assimilation percentage, %C : Consumption percentage

(5)

너지의

90.30%,

^총^{섭취에너지의}

21.53%

^인

17.24 J

^이었으며

,

이것은성장이

94.03%,

탈피체가

5.97%

를각각차지하였다

.

고 찰

해산동물의에너지흐름을파악하기위해서는사전에그생물 의생활사를정확하게조사하는것이필요하다

.

^갑각류의^성장 은환경에따라다양한반응을보이지만

(Hartnoll and Dalley,

1981),

^특히^섭취

,

^성장^및^대사율은^{생육단계별로도}^많은^변

동을보이는것으로알려져있다

(Reeve, 1969; Hiroko et al., 1990).

십각목갑각류유생의성장은일반적으로지수적성장을나 타낸다

(Mootz and Epifanio, 1974; Logan and Epifanio, 1978;

Johns and Pechenik, 1980; Stephenson and Knight, 1980;

Dawirs, 1983).

^본^연구에서^실내^사육한^{가시배새우의}^유생성 장은지수적성장을보였으며

, Park (1989)

^의^실험실^내에서^사 육한보리새우

, Penaeus japonicus

^의^{초기유생의}^성장에서^지 수적성장을보였다

.

그리고

Her (2002)

의물렁가시붉은새우

, Panadalopsis japonica

의초기유생에서의성장에서도 지수적 인성장을보임으로써

,

^다른^갑각류의^유생성장^양상과도^유사 하였다

(Dawirs, 1983; Jacobi and Anger, 1985; Harms, 1987;

Kim, 2000).

가시배새우 유생의 산소소비량은 체중이 유생발육이 진행 될수록증가하였다

.

^이는

Carcinus maenas (Dawirs, 1983)

와

Hyas coarctatus (Jacobi and Anger, 1985)

^의^연구된^결과에서 도다른갑각류의유생과유사한결과로이러한체중증가에따 른산소소비량의증가는갑각류의일반적인경향을나타내었 다

(Beamish, 1964; Fry, 1971; Dabrowski, 1986; Degani et al., 1989; Cai and Summerfelt, 1992).

수서무척추동물은동화에너지의거의

2/3

^를^호흡에^사용하는 것으로알려져있고

(Clutter and Theilacker, 1971),

^특히^생물 에있어서호흡을통한대사에너지는에너지수지구성요소별 사용에너지의백분율에많은변동을야기할수있다

. Chin et

al. (1992)

^은^호흡열^상실에^따른^에너지^계산은^동물이^서식

지에서경험하는수온범위내의여러수온구에서산소소비량 을측정하고

,

^이^{결과로부터}^얻어진^수온^및^체중과^상관관계 를갖는산소소비량회귀식을적용하는것이생물의호흡을통 한대사에너지에대한이해도를높일수있는방법이라고제안 하였다

.

가시배새우유생의섭취량은

Park (1989)

^이^보고한

Penaeus japonicus

의

zoea

^{유생보다는}^훨씬^높았으며

, mysis

^유생과^비 슷한섭취량으로나타났다

.

^그러나

Emmerson (1984)

^이^보고한

Penaeus indicus

의

protozoea

유생보다는훨씬낮게나타났다

.

본연구결과에서

23.84%

^의^{동화효율은}

, Penaeus japonicus

의

17.62% (Park, 1989)

^및

Carcinus maenas

^의

22.2% (Dawirs, 1983)

^와는 ^비슷한 ^수준을 ^보였다

.

^하지만

Rhithropanopeus harrisii

^의

45.0% (Levine and Sulkin, 1979), Menippe mer-

cenaria

^의

63.2% (Mootz and Epifanio, 1974)

^및

Cancer ir- roratus

의

77.0% (Johns, 1982), Neomysis intermedia

의암컷

85.65%,

^수컷

81.99% (Choi et al., 1997)

^에^비해서는^훨씬^낮 은값을보였다

.

이와같은낮은동화효율은종에따른차이도 있을수있지만

,

^환경수의^온도와^비례하는^{소화기능과}^밀접한 관련을가지고있기때문에

,

^본^{연구에서처럼}^한해성^새우의^경 우동화효율이낮은반면분으로배출되는에너지비율이높았 던것으로판단된다

.

가시배새우의 총 성장효율

(K

₁

)

은

21.53%

로

Penaeus ja- ponicus (Park, 1989)

및

Rhithopanopeus harrisii (Levine and Sulkin, 1979), Menippe mercenaria (Mootz and Epifanio, 1974)

와

Cancer irroratus (Johns, 1982)

의

23-39%

와유사한 결과를보였지만

, Carcinus maenas

의

3.3% (Dawirs, 1983), Neomysis intermedia

의암컷

8.71%,

수컷

9.02% (Choi et al., 1997), Pandalus hypsinotus

의

12.91% (Kim, 1994)

^보다^높 았다

.

^순 ^성장효율

(K

₂

)

^은

90.30%

^로

Pandalopsis japonica

^의

91.39% (Kim, 2000)

와비슷하게나타났으며동화량에대하여 성장으로전환된에너지가상당히높은것으로나타났다

.

^이러 한효율적차이는각각의종에대한유생단계와공급된먹이에 서많은영향을받는것으로판단된다

.

대사활동과정에서에너지의소비는에너지수지로구체적으 로설명된다

(Adams and Breck, 1990).

다양한생물학적

,

물리 적

,

^화학적^조건^하에서^오는^{에너지수지에}^관한^생체역학^모델 구축

(Bartell et al., 1986)

을위한일환에서본연구의가시배새 우유생에너지수지에대한구체적인정보는매우유용하게활 용될것으로판단된다

.

사 사

본 연구는농림축산식품부수산실용화기술개발사업에의해 이루어진것임

.

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