해양 생태계 내에서 혼합영양 섬모류 Myrionecta rubra의 중요성

명금옥,김형섭,장건강,박종우,이원호,Myung, Geum-Og,Kim, Hyung-Seop,Jang, Keon-Gang,Park, Jong-Woo,Yih, Won-Ho 한국해양학회 2007 바다 Vol.12 No.3

Myrionecta rubra Jankowski 1976(=Mesodinium rubrum Lohmann 1908)는 다양한 해양 환경에서 적조를 일으키는 매우 일반적인 원생생물인 혼합영양 섬모류이다. M. rubra의 실험실 종주가 처음 보고된 2000년 이후, M. rubra의 생태적 역할 및 진화적 중요성에 관한 새로운 논문들이 발표되고 있다. 본고는 M. rubra의 생태학적 중요성에 관한 새로운 연구 결과들을 중심으로, 이 혼합영양성 종에 대한 정확한 인식에 도움이 되는 종설로서 작성되었다. M. rubra는 Teleaulax sp.와 Geminigera sp. 등의 은편모류를 섭식한 후 이들의 색소체를 잔류시켜 광합성을 하는 "kleptoplastidic ciliate"이다. 최근에는 M. rubra의 박테리아 섭식이 확인되어, 지금까지 보고된 M. rubra의 영양유형은 광합성, 박테리아 섭식, 편모조류섭식 등으로 다양하다. M. rubra는 요각류, 곤쟁이류, 유즐동물과 멸치류의 유생, 그리고 이매패류 치패와 같은 metazoan predator의 먹이생물로 알려져 있다. 2006년에는 설사성 패독을 일으키는 원생생물인 와편모류 Dinophysis도 M. rubra를 섭식하는 섭식자 가운데 하나라는 사실이 새로이 발표되었다. 따라서, 해양 혼합영양 섬모류 M. rubra는 표영 생태계 먹이망(pelagic food web)의 핵심적 주요 linking ciliate의 하나로서 에너지 및 유기물 흐름에 크게 기여할 것이다. Myrionecta rubra Jankowski 1976(=Mesodinium rubrum Lohmann 1908), a mixotrophic ciliate, is very common and often causes recurrent red tides in diverse marine environments. Since the report on the first laboratory strain of this species in 2000, papers on its novel ecological role and evolutionary importance have been high lighted. This review paper is prepared to promote the de novo recognition M. rubra as a marine mixotrophic species. M. rubra is a ciliate which is able to photosynthesize using plastids originated from cryptophyte (including Teleaulax sp. and Geminigera sp.) prey cells (i.e. kleptoplastidic ciliate). Recently, novel bacterivory of M. rubra was firstly reported. Thus, the nutritional modes of M. rubra include photosynthesis, bacterivory, and algivory. In turn, M. rubra was reported as the prey species of metazoan predators such as calanoid copepods, mysids, larvae of ctenophore and anchovy, and spats of bivalves. In addition, it was reported that dinoflagellate Dinophysis causing diarrhetic shellfish poisoning is one among the predators of M. rubra. Thus, M. rubra, a marine mixotrophic ciliate, may play a pivotal role as a common linking ciliate for the flow of energy and organic material in pelagic food webs.

광적응된 은편모조류를 섭식한 해양 광합성 섬모류 Mesodinium rubrum MR-MAL01의 흡광스펙트럼

김형섭,명금옥,조수근,이원호 한국해양학회 2003 바다 Vol.8 No.1

한국 서해의 곰소만에서 분리한 온대 해역 산 초유의 M. rubrum MR-MAL01 배양체가 확보되어, M. rubrum의 입자성 먹이 섭식 현상이 보고되었다. 100$\mu$mo1 photons m$^{-2}$ s$^{-1}$의 고광도(HL) 및 10 $\mu$mo1 photons m$^{-2}$ s$^{-1}$의 저광도(LL)에서 광 적응된 미동정 은편모조류 종주 CR-MAL01를 섭식한 M. rubrum의 HL및 LL적응 배양체의 흡광스펙트림을 분석하였다. LL에 적응된 은편모조류를 섭식하고 LL에 적응한 M. rubrum은 섭식 이전의 먹이 세포 흡광특성인 542nm부근의 황갈색 파장 범위에서 뚜렷한 흡광피크를 나타낸 반면, HL에 적응된 은편모조류를 섭식하고 HL에 적응한 M. rubrum은 이러한 흡광 피크를 보이지 않았다. 이는 M. rubrum세포가 자연환경에서 먹이로서 은편모조류 세포를 섭식하기 이전 이들의 광 적응 상태가 섭식 이후 M. rubrum의 광이용 효율 및 광흡수 특성 등을 결정짓는 주요 요인이 될 가능성을 시사한다. 본 연구 결과에 따라 예상되는 해양 미소생물 생태계 내에서 이들 섭식자 및 먹이생물 종의 역할에 대하여 논의하였다. Recent reports on the phagotrophic feeding of M. rubrum are based on cultivation experiments with novel isolates of this ciliate species from Gomso Bay, Korea. Photo-adapted cryptophyte(CR-MAL01) cultures at high light of 100 $\mu$mol photons m$^{-2}$ s$^{-1}$ (HL) and low light of 10 $\mu$mol photons m$^{-2}$ s$^{-1}$ (LL) were fed to M. rubrum (MR-MAL01) cultures under HL and LL conditions, respectively. Absorbance spectrum by LL M. rubrum showed the same peak at wavelengths around 542nm as that by LL cryptophyte prey, which was not showed in HL M. rubrum. This result supports the implication that light utilization and absorption pattern of M. rubrum population must depend on the status of photo-adaptation of the co-existing population of prey cryptophyte. Consequences of the present research results were discussed in relation to the function of the prey cryptophyte and phagotrophic M. rubrum in marine microbial ecosystem.

춘계 금강 하구에서 혼합영양 섬모류인 Myrionecta rubra (=Mesodinium rubrum) 개체군의 단주기 변동

이원호,명금옥,김형섭,정해진,Yih, Won-Ho,Myung, Geum-Og,Kim, Hyung-Seop,Jeong, Hae-Jin 한국조류학회(藻類) 2005 ALGAE Vol.20 No.3

Myrionecta rubra, a mixotrophic ciliate, is a cosmopolitan red tide species which is commonly found in neritic and estuarine waters. M. rubra had long been listed as an “nculturable protist”until 2 different laboratory strains were finally established in 2 research groups at the beginning of this century, enabling us to perform initiative investigation into various aspect of the live M. rubra strains (Gustafson et al. 2000; Yih et al. 2004b; Johnson and Stoecker 2005). Field sampling was carried out on high tide at 2 fixed stations around Kunsan Inner Harbor (St.1 near the Estuarine Weir and St.2 off Kunsan Ferry Station) every other day for 4 months from mid-February 2004 to understand detailed figure of the recurrent spring blooms of M. rubra following the onset of the water gates operation of the Keum River Estuarine Weir on August 1994. With its maximum abundance of 272 cells mL$^{-1}$ in St.1, fluctuation pattern of the M. rubra population at the 2 stations was strikingly similar. Notable growth of M. rubra population started on late April, to cause M. rubra red tides during one month from mid-May in which “xceptionally low salinity days”without its red tide were intermittently inserted. High abundance of M. rubra over 50 cells mL$^{-1}$ was recorded at samples with their water temperature and salinity higher than 15${^{\circ}C}$ and 4.0 psu, respectively. During pre-bloom period when salinity fluctuation is moderate and the water temperature is cooler than 15°C, Skeletonema costatum, a chain-forming centric diatom, was most dominant. Cyanobacterial species such as Aphanizomenon flos-aquae and Phormidium sp. replaced other dominant phytoplankters on the days with “xceptionally low salinity”even during the main blooming period of M. rubra. To summarize, M. rubra could form spring blooms in Keum River Estuary when the level of salinity fluctuation was more severe than that for the dominant diatom Skeletonema costatum and milder than that for the predominance by freshwater cyanobacteria. Therefore, optimal control of the scale and frequency of freshwater discharges might lead us to partially modify the fluctuation pattern of M. rubra populations as well as the period of spring blooms by M. rubra in Keum River Estuary. Sampling time interval of 2 days for the present study or daily sampling was concluded to be minimally required for the detailed exploration into the spring blooms by M. rubra populations in estuaries with weirs like Keum River Estuary.

금강하구언 담수방류와 춘계 식물플랑크톤 군집의 단주기 변동

이원호,명금옥,유영두,김영길,정해진,Yih, Won-Ho,Myung, Geum-Og,Yoo, Yeong-Du,Kim, Young-Geel,Jeong, Hae-Jm 한국해양학회 2005 바다 Vol.10 No.3

1988년에 완공하여 1994년 8월부터 수문을 작동한 금강 하구언으로 인해 금강 하구역 환경은 크게 변하였다. 수문을 통해 담수를 불규칙하게 방류하면, 극심한 염분 저하와 함께 수중 무기영양염 농도가 급변한다. 이와 같이 환경 스트레스가 큰 금강 하구 내측에서 춘계 식물플랑크톤 군집의 단주기 변동을 연구하기 위하여, 2004년 2월 중순부터 4개월 동안 2개의 고정점에서 약 2일 간격으로 고조 때에 현장조사와 시료채취를 실시하였다. 하구언에 근접한 정점에서는 정점2(도선장 부근 정점)에 비해 염분 측정치의 변동계수가 2배에 이르러, 정점1에서 시간에 따른 염분의 변동이 극심함을 반증하였다. 용존 무기영양염 가운데, 질소계 영양염($NO_3^-,\;NO_2^-$ 및 $NH_4^+$)의 농도가 정점2에 비해 정점1에서 뚜렷이 높게 나타나, 수문 작동에 따라 그 농도가 부정기적으로 급변할 수 있는 환경임을 확인하였다. 식물플랑크톤 군집을 구성하는 주요 분류군은 규조류로서 출현종의 수나 생물량 모두 최고치를 기록하였다. 이 가운데 단독성 중심 규조 Cyolotella meneghiniana와 사슬형 중심 규조 Skeletonema costatum은 조사기간 전반부에 차례대로 5-6주 동안씩(천이 구간 I 및 II)절대 우점하여, 수온 $10^{\circ}C$의 시점을 중심으로 뚜렷한 천이현상을 보여 주었다. 천이구간 III(4월 중순-5월 말)은 대량 방류된 담수와 함께 하구역에 유입된 담수 녹조종인 남세균 Aphanizomenon flos-aquae와 Phormidium sp. 등이 매우 높은 구성비를 나타낸 시기이다. 이 기간에는 극저염-고염의 극심한 변동으로 인하여 낮은 수준의 생물량을 기록한 규조 Cyclotella meneghiniana와 Skeletonema costatum 외에는 담수종들이 전적으로 우점하였다. 천이구간 IV에서는 수온이 $18^{\circ}C$ 이상으로 상승하면서, 규조 Guinardia delicatuia가 단일 시료 내 최고 우점도 $75^{\circ}C$를 기록하며 약 일주일간 우점하였다. 즉, 조사기간 중에는 수온 $18^{\circ}C$ 미만의 극심한 염분 변동기(천이구간 III)를 제외한, 전 기간 중에 규조가 절대 우점하며 춘계 식물플랑크톤 우점종의 천이를 주도하였다. 천이구간 III에서도, 만약 담수방류의 규모와 빈도를 적절히 조절하였더라면, 주요 우점종이 담수 남세균이 아닌 하구성 규조로 바뀔 수도 있었을 것이다. 본 연구에 적용한 격일간 현장조사는 불규칙한 담수방류가 불가피한 반일주조 하의 하구역 환경에서 식물플랑크톤 종의 천이현상을 규명하기 위한 최소한의 필요조건으로 판단된다. Irregular discharges of freshwater through the water gates of the Keum River Estuarine Weir, Korea, whose construction had been completed in 1998 with its water gates being operated as late as August 1994, drastically modified the estuarine environment. Sharp decrease of salinity along with the altered concentrations of inorganic nutrients are accompanied with the irregular discharges of freshwater into the estuary under the influence of regular semi-diurnal tidal effect. Field sampling was carried out on the time of high tide at 2 fixed stations(St.1 near the Estuarine Weir and St.2 off Kunsan Ferry Station) every other day for 4 months from mid-February 2004 to investigate into the semi-weekly variation of spring phytoplankton community in relation to the freshwater discharges from Keum River Estuarine Weir. CV(coefficient of variation) of salinity measurements was roughly 2 times greater in St.1 than that in St.2, reflecting extreme salinity variation in St.1 Among inorganic nutrients, concentrations of N-nutrients($NO_3^-,\;NO_2^-$ and $NH_4^+$) were clearly higher in St.1, to imply the more drastic changes of the nutrient concentrations in St.1. than St.2 following the freshwater discharges. As a component of phytoplankton community, diatoms were among the top dominants in terms of species richness as well as biomass. Solitary centric diatom, Cyclotella meneghiniana, and chain-forming centric diatom, Skeletonema costatum, dominated over the phytoplankton community in order for S-6 weeks each (Succession Interval I and II), and the latter succeeded to the former from the time of <$10^{\circ}C$ of water temperature. Cyanobacterial species, Aphanizomenon Posaquae and Phormidium sp., which might be transported into the estuary along with the discharged freshwater, occupied high portion of total biomass during Succession Interval III(mid-April to late-May). During this period, freshwater species exclusively dominated over the phytoplankton community except the low concentrations of the co-occurring 2 estuarine diatoms, Cyclotella meneghiniana and Skeletonema costatum. During the 4th Succession Interval when the water temperature was over $18^{\circ}C$, the diatom, Guinardia delicatula, was predominant for a week with the highest dominance of $75\%$ in discrete samples. To summarize, during all the Succession Intervals other than Succession Interval III characterized by the extreme variation of salinity under cooler water temperature than $18^{\circ}C$, the diatoms were the most important dominants for species succession in spring. If the scale and frequency of the freshwater discharge could have been adjusted properly even during the Succession Interval III, the dominant species would quite possibly be replaced by other estuarine diatom species rather than the two freshwater cyanobacteria, Aphanizomenon flosaquae and Phormidium sp.. The scheme of field sampling every other day for the present study was concluded to be the minimal requirement in order to adequately explore the phytoplankton succession in such estuarine environment as in Keum River Estuary: which is stressed by the unpredictable and unavoidable discharges of freshwater under the regular semi-diurnal tide.

국내 연안산 질소고정 단세포 남세균 종주의 광생물학적 수소생산력

박종우,명금옥,이원호 한국수소및신에너지학회 2010 한국수소 및 신에너지학회논문집 Vol.21 No.2

Photobiological hydrogen production by nitrogen-fixing unicellular cyanobacteria has long been considered to be an environmentally sound and very promising method for the future supply of renewable clean energy. We tried to find out the optimum cell concentration for H2 production in each of the two new Korean nitrogen-fixing unicellular cyanobacterial strains to compare with Synechococcus sp. strain Miami BG043511. The two Korean strains, Cyanothece sp. KNU CB MAL-031 and KNU CB MAL-058, were isolated from Korean west coasts. Cell concentrations up to 17 billion cells ml-1 were applied to the tests. High cell concentration over 15 billion cells ml-1 resulted in drastically reduced H2 production in all the three strains. The two domestic strains, however, produced 2-3 time more hydrogen than Synechococcus sp. Miami BG043511 at cell concentrations of 5-10 billion cells ml-1. At lower cell concentrations than 2 billion cells ml-1, MAL-031 exhibited highest H2 production followed by Miami BG043511, with far less production in MAL-058. Present result suggests that Cyanothece sp. MAL-CB031 might be one of the ideal nitrogen-fixing unicellular cyanobacterial strains for the photobiological hydrogen production.

Ingestion rate and grazing impact by the mixotrophic ciliate Mesodinium rubrum on natural populations of marine heterotrophic bacteria in the coastal waters of Korea

성경아,명금옥,정해진,이원호,김형섭,Hyun Jung Jo,박재연,유영두 한국조류학회I 2017 ALGAE Vol.32 No.1

We explored feeding by the mixotrophic ciliate Mesodinium rubrum, heterotrophic nanoflagellates (HNFs), and small ciliates (<30 μm in cell length) on natural populations of heterotrophic bacteria in Masan Bay, Keum River Estuary, and in the coastal waters of the Saemankeum area, Korea when M. rubrum red tides occurred. We also measured ingestion rates of M. rubrum on cultured heterotrophic bacteria as a function of bacterial concentration in the laboratory. The ingestion rates of M. rubrum on natural populations of heterotrophic bacteria (2.3–16.8 bacteria grazer−1 h−1) were comparable to or lower than those of co-occurring HNFs (10.7–41.7 bacteria grazer−1 h−1), but much lower than those of co-occurring small ciliates (76.0–462.2 bacteria grazer−1 h−1). However, the maximum grazing coefficient of M. rubrum (0.245 d−1) on natural populations of heterotrophic bacteria was much higher than that of small ciliates (0.089 d−1), and slightly higher than that of HNFs (0.204 d−1). With increasing bacterial concentrations, ingestion rates of M. rubrum on cultured heterotrophic bacteria continuously increased, but became saturated at higher prey concentrations over 1–5 × 106 cells mL−1. The maximum ingestion rate of M. rubrum on cultured heterotrophic bacteria was 34.4 bacteria grazer−1 h−1. Based on the present study, it is suggested that M. rubrum may be an important grazer of heterotrophic bacteria and sometimes have considerable grazing impact on natural populations of heterotrophic bacteria.

Optimal growth conditions for the two euryhaline cyanobacterial clones, Anabaena spp. CB-MAL21 and CB-MAL22 isolated from Mankyeong Estuary, Korea

김영길,명금옥,이원호,신윤근 한국조류학회I 2004 ALGAE Vol.19 No.2

As a result of the 2-year monthly monitoring of the phytoplankton community at 3 stations in Mankyeong Estuary, Korea, we learned that cyanobacterial species of the genus Anabaena occurred at most sampling points with huge salinity differences (0.1-32.5 psu). We isolated several clones of Anabaena spp. from the monitoring stations, and screen out two euryhaline and nitrogen-fixing Anabaena clones, CB-MAL21 and CB-MAL22. The two clones were grown under various environmental gradients such as temperature (20, 30, 35 and 40°C), salinity (0, 2, 5, 15 and 30 psu), and PO4 3--P concentration (0, 1.6, 8.0, 40 and 200 μM). Growth of CB-MAL21 and CB-MAL22 was measured by daily monitoring of chlorophyll fluorescence from each experimental culture for more than three serial transfers. Both the two experimental clones did not grow at 0 psu. Maximal growth rates of the two clones were markedly reduced at lower PO4 3--P concentrations showing negligible growth at 0 and 1.6 μM. However, growth of CB-MAL21 was not affected by low NO3 --N concentration in culture media, showing the nitrogen-fixing ability. Maximum biomass yields of the two clones decreased dramatically at 35 and 40°C. Optimal growth conditions for the two experimental clones were determined to be 20-30°C, 40 μM PO4 3--P, and wide salinity range from 5.0 to over 30 psu. Best growth of CB-MAL21 was shown at (20°C-15 psu), which is less saline and cooler condition than those (i.e., 30°C-30 psu) for the best growth of CB-MAL22. The euryhaline and nitrogen-fixing CB-MAL21 strain thus can be a candidate laboratory culture for the future cyanobacterial marine biotechnology in temperate coastal waters.

Ingestion of the unicellular cyanobacterium Synechococcus by the mixotrophic red tide ciliate Mesodinium rubrum

유영두,성경아,명금옥,김형섭,정해진,Brian Palenik,이원호 한국조류학회I 2015 ALGAE Vol.30 No.4

We explored phagotrophy of the phototrophic ciliate Mesodinium rubrum on the cyanobacterium Synechococcus. The ingestion and clearance rates of M. rubrum on Synechococcus as a function of prey concentration were measured. In addition, we calculated grazing coefficients by combining the field data on abundance of M. rubrum and co-occurring Synechococcus spp. with laboratory data on ingestion rates. The ingestion rate of M. rubrum on Synechococcus sp. linearly increased with increasing prey concentrations up to approximately 1.9 × 106 cells mL-1, to exhibit sigmoidal saturation at higher concentrations. The maximum ingestion and clearance rates of M. rubrum on Synechococcus were 2.1 cells predator-1 h-1 and 4.2 nL predator-1 h-1, respectively. The calculated grazing coefficients attributable to M. rubrum on cooccurring Synechococcus spp. reached 0.04 day-1. M. rubrum could thus sometimes be an effective protistan grazer of Synechococcus in marine planktonic food webs. M. rubrum might also be able to form recurrent and massive blooms in diverse marine environments supported by the unique and complex mixotrophic arrays including phagotrphy on hetrotrophic bacteria and Synechococcus as well as digestion, kleptoplastidy and karyoklepty after the ingestion of cryptophyte prey.

광합성 섬모류 Mesodinium rubrum MR-MAL01의 응용성 (1) 이매패류 종묘생산을 위한 먹이생물 유용성

김형섭(Hyung-Seop Kim),명금옥(Geumog Myung),조수근(Soo-Gun Jo),이원호(Wonho Yih) 한국수산과학회 양식분과 2004 韓國養殖學會誌 Vol.17 No.2

Abundance of Epiphytic Dinoflagellates from Coastal Waters off Jeju Island, Korea During Autumn 2009

김형섭,이원호,김종혁,명금옥,정해진 한국해양과학기술원 2011 Ocean science journal Vol.46 No.3

The occurrence of harmful epiphytic dinoflagellates is of concern to scientists, the aquaculture industry, and government due to their toxicity not only to marine organisms but also to humans. There have been no studies on the abundance of the epiphytic dinoflagellates in Korean waters. We explored the presence of epiphytic dinoflagellates in the coastal waters off Jeju Island, southwestern Korea. Furthermore,we measured the abundance of epiphytic dinoflagellates on the thalli of 24 different macroalgae, collected from five different locations in October 2009. Five epiphytic dinoflagellate genera Amphidinium, Coolia, Gambierdiscus, Ostreopsis, and Prorocentrum were found. These five genera were observed on the thalli of the macroalgae Chordaria flagelliformis, Martensia sp., Padina arborescens, and Sargassum sp., while none were observed exceptionally on Codium fragile. The abundance of Ostreopsis spp. was highest on Derbesia sp. (8,660 cells/g wet weight),while that of Gambierdiscus spp. was highest on Martensia sp. (4,870 cells/g-ww). The maximum abundances of Amphidinium spp., Coolia spp., and Prorocentrum spp. were 410, 710, and 300 cells/g-ww, respectively. The maximum abundance of Coolia spp., Gambierdiscus spp., and Ostreopsis spp. obtained in the present study was lower than for other locations reported in literature. The results of the present study suggest that the presence and abundance of epiphytic dinoflagellates may be related to the macroalgal species of the coastal waters of Jeju Island.