Feeding and egg production of the planktonic copepod Calanus sinicus in spring and autumn in the Yellow Sea, China

Shipboard incubations were conducted in spring (April) and autumn (October/November) 2006 to measure the feeding and egg production rates (EPR) of Calanus sinicus in the Yellow Sea, China. The ingestion rate (2.08-11.46 and 0.26-3.70 mu g C female(-1) day(-1) in spring and autumn, respectively) was positively correlated with microplankton carbon concentrations. In the northern part of the Yellow Sea, feeding on microplankton easily covers the respiratory and production requirements, whereas in the southern part in spring and in the frontal zone in autumn, C. sinicus must ingest alternative food sources. Low ingestion rates, no egg production and the dominance of the fifth copepodite (CV) stage indicated that C. sinicus was in quiescence inside the Yellow Sea Cold Bottom Water (YSCBW) area in autumn. Calanus sinicus ingested ciliates preferentially over other components of the microplankton. The EPR (0.16-12.6 eggs female(-1) day(-1) in spring and 11.4 eggs female(-1) day(-1) at only one station in autumn) increased with ciliate standing stock. Gross growth efficiency (GGE) was 13.4% (3-39%) in spring, which was correlated with the proportion of ciliates in the diet. These results indicate that ciliates have higher nutrient quality than other food items, but the low GGE indicates that the diet of C. sinicus is nutritionally incomplete.

Effects of single and mixed diatom diets on the reproduction of copepod Calanus sinicus

Under laboratory conditions, the potential influence of diatom diets on reproduction of zoo-plankton Calanus sinicus was studied. Four diatom diet ingredients: Skeletonema costatum (SC), Chaetoceros muelleri (CM), Phaeodactylum tricornutum (PT), diatom mixture (MIX) and a control diet: the flagellate Platymonas subordiformis (PS), were used at the same carbon concentrations of 2.0 mu g mL(-1) C. In a period of 17-day laboratory experiment, the effects of these algae diets on egg production and hatching success of the copepod Calanus sinicus were examined. The diets were analyzed for fatty acid content as an indicator of food quality. The results showed that the female survival of all treatments reached more than 80% except PT. Comparing to the initial value, egg production of Calanus sinicus was reduced in diatom diets (PT, CM), but remained in normal level in SC and MIX, indicating that some single diatom diets had a negative effect on the egg production of Calanus sinicus. Feeding with mixed food however can eliminate this negative effect. Among all the treatments, hatching success in filtered seawater was significantly higher than in algal exudates, indicating that not only diatoms but also other phytoplankton in certain concentration can release extracelluar substance that may inhibit eggs from hatching. Fatty acid analysis showed that both egg production rate and hatching success were negatively correlated to the ratio of 20:5 omega 3 and 14:0 in fatty acid composition.

The combined effects of temperature and food supply on Calanus sinicus in the southern Yellow Sea in summer

The effects of temperature and food availability on the life history strategy of the planktonic copepod Calanus sinicus in the southern Yellow Sea in summer were studied in this paper. The fifth copepodite stage (CV) dominates the population in the central part of the southern Yellow Sea, where the Yellow Sea Cold Water Mass (YSCWM) occurs below the thermocline. Incubation experiments were conducted on CV C. sinicus caught from the YSCWM to examine the effects of temperature and food availability. Temperature at the surface (27degreesC) is lethal to CVs regardless of food availability. At the temperature in the middle of the thermocline (18degreesC), survival time of the specimens depends on food availability, being similar to20 days in treatments without extra food supply. At the temperature in the YSCWM (9degreesC), most animals survive at the end of 27 day incubation even in treatments without food supply. Developmental rate of CVs at 9degreesC without extra food supply is extremely low. The increase of either temperature or food supply promotes the developmental rate of CVs. According to these results, the surface layers with high temperature and low food abundance are detrimental for the survival and reproduction of C. sinicus. Low temperature and low food availability in the YSCWM help CV to maintain a much lower developmental rate and higher survival rate. The ecological trait of C. sinicus in the southern Yellow Sea in summer cannot be sufficiently explained solely by the effects of temperature.

Life history strategies of Calanus sinicus in the southern Yellow Sea in summer

Ecological and physiological features of the planktonic copepod Calanus sinicus in the southern Yellow Sea in summer were studied to reveal its life history strategy. From the coastal shallow waters to the central part of the southern Yellow Sea, a shift of the stage composition occurs from being dominated by the egg-nauplius stage to being dominated by the fifth copepodite (CV) stage. Most CVs reside in the Yellow Sea Cold Water Mass (YSCWM), where both temperature and food abundance are low. CVs in the YSCWM have longer body lengths, heavier body weights and higher carbon contents than those outside the YSCWM. Onboard incubations show that the development of CVs in the YSCWM is suspended. Energy conservation, development suspension and lack of diel vertical migration (DVM) behavior suggest a diapause status for the CVs in the YSCWM, although vertical distribution patterns indicate the CV individuals are not fully synchronous in physiology and development. This adaptive oversummering strategy would help C. sinicus to live through the warm and food-limited summer in the central part of the southern Yellow Sea; both low temperature and low food supply are necessary for CV to maintain the resting state in the YSCWM. Calanus sinicus exhibits different life history strategies in different regions of the southern Yellow Sea in summer.

Summer reproduction of the planktonic copepod Calanus sinicus in the Yellow Sea: influences of high surface temperature and cold bottom water

We have observed that Calanus sinicus retreated from neritic areas in the Yellow Sea and concentrated in the Yellow Sea Cold Bottom Water (YSCBW) area in summer. To investigate the summer reproductive strategy of C. sinicus in this situation, effects of high temperature on reproduction and hatching, as well as geographical variation of in situ egg production rate, were studied by onboard incubation in August 2001. Diel vertical migration (DVM) of females was investigated within and outside the YSCBW, respectively. Onboard incubation at 27 degrees C (i.e. surface temperature) resulted in lower fecundities than that at 9.8 and 12 degrees C (i.e. bottom temperature inside and outside the YSCBW) together with decreased hatching rates and increased naupliar malformation. Egg production was more active at stations outside the YSCBW than inside, where chlorophyll-a concentration was also relatively low. Females inside the YSCBW underwent DVM although they rarely entered the surface layer, but DVM was not observed outside the YSCBW. We conclude that surface temperature in summer has deleterious effects on C. sinicus egg production and hatching, and that it cannot reproduce successfully over the whole area. Inside the YSCBW, egg production is depressed by low food availability, while females outside suffer from high temperatures because of strong vertical mixing.

Seasonal variation of reproduction rates and body size of Calanus sinicus in the Southern Yellow Sea, China

Annual variations of egg production rate (EPR) and clutch size of Calanus sinicus, as well as body size of females (prosome length and dry weight), were investigated at a series of stations in the Southern Yellow Sea by onboard incubation. Calanus sinicus was spawning in all the 11 cruises investigated, and the annual variation of EPR was bimodal. Monthly average EPR was highest from May to July, respectively, 5.97, 5.36 and 6.30 eggs female(-1) d(-1), then decreased dramatically to only 1.37 eggs female(-1) d(-1) in August and attained the lowest 1.07 eggs female(-1) d(-1) in October. In November, average EPR increased again to 4.31 eggs female(-1) d(-1). Seasonal variation of clutch size was similar to EPR, except that it decreased gradually after August rather than dramatically as did EPR. Prosome length of females was maximum in May and minimum in October, but dry weight was highest in November. Monthly average EPR correlated better with prosome length than dry weight, while clutch size was rather determined by dry weight of females. It is suggested that egg production of C. sinicus was active during two discontinuous periods when both surface and bottom temperature fell into its favorite range (i.e. 10-23degreesC), and different reproductive strategies were adopted in these two reproductive peaks: other than the highest EPR, longer prosome length was also achieved by C. sinicus from May to July, while females in November developed shorter bodies but accumulated more energy for reproduction.

Spring and autumn reproduction of Calanus sinicus in the Yellow Sea

The fecundity, hatching success and naupliar survival of Calanus sinicus were studied in the Yellow Sea during research cruises in April and October 2006, with emphasis on the regulation of reproduction. During both cruises, the egg production rate (EPR) showed large spatial variations (0 to 25.4 eggs female(-1) d(-1)), generally coinciding with the food availability. In April, the abundant phytoplankton and ciliates in the study area supported active reproduction, which would probably initiate the annual population development. In October, females remained immature in the Yellow Sea Cold Water Mass, likely due to the unfavorable environment (poor food and low temperature). However, reproduction and recruitment were high in the neritic region, which may explain the local population recovery in late autumn. Hatching success varied markedly among stations in April (4 to 85%), whereas it was high overall in the neritic region during October (> 90%). Based on the potential recruitment rates, the spring recruitment would be more important for the annual population dynamics, Female gonad maturity, body size and lipid reserve were examined in relation to fecundity. Regression analyses suggested that the reproductive index (defined as the proportion of females with mature gonads) could be an indicator for the EPR of C. sinicus. Among the regulating factors, external food (ciliates and phytoplankton) seems essential for reproduction, whereas inner lipid reserve may mainly serve metabolic needs. Moreover, fecundity is positively related to body size but independent of temperature, which might exert indirect influences on reproduction.

Feeding and respiration rates of a planktonic copepod (Calanus sinicus) oversummering in Yellow Sea Cold Bottom Waters

The distribution, feeding and oxygen consumption of Calanus sinicus were studied in August 2001 on a transect across Yellow Sea Cold Bottom Waters (YSCBW) and two additional transects nearby. The distribution of C. sinicus adults and copepodites stage CV appeared to be well correlated with water temperature. They tended to concentrate in the YSCBW (>10,000 ind. m(-2)) to avoid high surface temperature. Gut pigment contents varied from 0.44 to 2.53 ng chlorophyll a equivalents (chl a equiv.) ind.(-1) for adults, and from 0.24 to 2.24 ng chl a equiv. ind.(-1) for CV copepodites. We found no relationship between gut pigment contents and the ambient chl a concentrations. Although the gut evacuation rate constants are consistent with those measured for other copepods, their low gut pigment contents meant an estimated daily herbivorous ingestion of <3% of body carbon in the YSCBW and <10% outside the YSCBW. However, based on estimates of clearance rates, C. sinicus feeds actively whether in the YSCBW or not, so the low ingestion rates probably reflect shortage of food. Oxygen consumption rates of C. sinicus ranged from 0.21 to 0.84 mul O-2 ind.(-1) h(-1), with high rates often associated with high temperature. From the oxygen consumption rates, daily loss of body carbon was estimated to be 4.0-13.7%, which exceeds our estimates of their carbon ingestion rates. C. sinicus was probably not in diapause, either within or outside the YSCBW, but this cold-water layer provides C. sinicus with a refuge to live through the hot, low-food summer.

Abundance of Calanus sinicus across the tidal front in the Yellow Sea, China

The abundance of Calanus sinicus eggs, nauplii, copepodites and adults and chlorophyll a (Chl-a) concentration were studied across tidal fronts in October 2000, and May and June 2001 in the Yellow Sea, China. The aim of the study was to evaluate the role of tidal fronts in the ecology of C. sinicus. The hydrographic tidal fronts were identified by the horizontal temperature gradient in the bottom layer and the temperature profiles across the fronts. The survey results showed that the concentration of Chl-a was high in the vicinity of the fronts, particularly in spring and early summer. The abundance of C. sinicus eggs and nauplii was usually higher in the tidal fronts than in the adjacent areas. In May and June 2001, the abundance of copepodites and adults of C. sinicus peaked in the tidal front. In June 2001 and October 2000, many copepodites and adults were found in stratified region.

The Yellow Sea cold bottom water - an oversummering site for Calanus sinicus (Copepoda, Crustacea)

The vertical distribution and stage-specific abundance of Calanus sinicus were investigated on three key transects in the southern Yellow Sea and the northern East China Sea in August 1999. The results showed that in summer C. sinicus shrank its distribution area to the central cold (less than or equal to10degreesC) bottom water in the Yellow Sea, i.e. the Yellow Sea Cold Bottom Water, remaining in high abundance (345.7 ind m(-3)). In the northern East China Sea on a transect from the mouth of the Yangtze River to the Okinawa trench, only a few individuals appeared in the inner side and none had been found either in the upper layer or in the deep layer of the outer shelf area. The population of C. sinicus in YSCBW consisted of mainly adults (46.83%) and C5 (37.41%). C1-C4 only accounted for 15.76%. The low proportion of the earlier copepodite stages and the high female:male ratio (11.39) indicated that the reproduction of C. sinicus in YSCBW was at a very low level due to the low temperature and low food concentration. It is concluded that the dramatic decrease of C. sinicus population in the shelf area of China seas in summer is caused by the shrinkage of its distribution area and the YSCBW served as an oversummering site.

Digestive gut structure and activity of protease, amylase, and alkaline phosphatase in Calanus sinicus during summer in the Yellow Sea and the East China Sea

Calanus sinicus aggregate at the depth of 40-60 m (ambient temperature is 16 degreesC) in the waters of the continental shelf of the Yellow Sea during summer. in animals found in near shore regions, there are changes in digestive gut cells structure, digestive enzyme activity (protease, amylase), and tissue enzyme (alkaline phosphatase (ALP)), which may represent adaptations by this cold-water animal to a sharp seasonal increase in temperature of 6-23 degreesC. The activities of the digestive enzymes (protease and amylase) are very low in animals at stations near the estuary of Yangtse River, whereas they are relatively high in animals at stations in the central Yellow Sea, During summer, B-cells of the intestine and the villi intestinalis disappear in animals that do not feed at stations near the estuary of the Yangtse River. Respiration rates were undetectable or quite low during summer in C. sinicus from stations near the estuary of the Yangtse River, whereas they were relatively high at stations in the central Yellow Sea. Based upon the morphological characteristics of the digestive gut structure, enzyme levels, respiration rates, and the distribution of C. sinicus, we concluded that C. sinicus might be dormant during summer in the near shore areas of the East China Sea while remaining active in the central Yellow Sea. (C) 2002 Elsevier Science B.V. All rights reserved.

Impact of microzooplankton and Calanus sinicus (Brodsky, 1962) on phytoplankton in the Yellow Sea during early summer

Dilution incubations and Calanus sinicus addition incubations were simultaneously conducted at five stations in the Yellow Sea in June of 2004 to evaluate the impact of microzooplankton and Calanus sinicus on phytoplankton based on the Chlorophyll a (Chl-a) levels. The Chl-a growth rates (k) ranged from 0.60-1.67 d(-1), while microzooplankton grazed the Chl-a at rates (g) of 0.29-0.62 d(t-1). The addition of C. sinicus enhanced the Chl-a growth rate (Z) by 0.004-0.037 d(-1) ind.(-1) L. C. sinicus abundance ranged from 84.1-160.9 ind. m(-3), which occupied 90.7%-99.1% of the copepod (> 500 mu m) population. The in-situ increase in phytoplankton by C. sinicus community was estimated to be 0.000 4-0.005 9 d(-1). These results showed that microzooplankton were the main grazers of phytoplankton, while C. sinicus induced a slight increase in the levels of phytoplankton.

Feeding habits of Calanus sinicus (Crustacea : Copepoda) during spring and autumn in the Bohai Sea studied with the herbivore index

Pigment ingestion rate (PIR) and egg production rate (EPR) of the dominant copepod Calanus sinicus, as well as chlorophyll-a concentration and phytoplankton assemblages were measured in the Bohai Sea, North China in June 1997, October 1998 and May 1999. A herbivore index (H) was also calculated as the carbon specific ratio of PIR and EPR, in order to investigate its feeding habits in the spring and autumn phytoplankton bloom respectively. On average, chlorophyll-a concentration was relatively similar (1-1.34 mg m(-3)) in the three cruises, but PIR was quite different. It was 3.24 mu g C female(-1) d(-1) in October, equivalent to one half of the PIR for June and one third of the PIR for May. Average EPR was highest in May, and quite similar during the other two months. According to H values, herbivorous feeding contributed 100% of the egg production of C. sinicus in June, 82.5% in May, but only 47.8% in October. It is possible that omnivorous feeding of C. sinicus in October was induced by a prevalence of large-sized diatoms and sufficient non-phytoplankton food resources during the autumn bloom period.

胶州湾四种哲水蚤种群动态的研究

本文研究了胶州湾内1988年5月至1989年5月，小拟哲水蚤[Paracalanus parvus(Claus)]、双刺纺锤水蚤[Acartia bifilosa (Giesbrecht)]、太平洋纺锤水蚤(A. pacifica steuer)和1982年5月至1983年4月中华哲水蚤(Calanus sinicus Brodsky)的种群结构和种群数量的季节变化。双剌纺锤水蚤和太平纺锤水蚤在胶州湾是季节性出现的种类，它们能产生休眠卵，本文对其休眠卵数量的季节变化以及影响休眠卵产生和孵化的因素进行了初步的探讨。本文还研究了体长的季节变化以及温度和食物条件对体长的影响。

长江口北港和北支浮游动物群落生态学比较研究

浮游动物是河口生态系统的重要组成部分，在食物网中，浮游动物是浮游植物捕食者，对浮游植物有摄食压力，所以控制浮游植物数量的增长。同时，浮游动物还是鱼类的饵料，从而影响渔业资源的数量。长江口是各种经济鱼、虾和蟹的重要产卵场、索饵场和洄游场。长江口各个分汊的分流量的差异较大，导致不同分汊的水文环境明显不同，其中最为明显就是北港和北支水域。然而以往对浮游动物的研究仅限于某一水域，没有对整体水域进行研究，本文选择最具代表性的北港和北支水域，对两水域浮游动物的群落生态特征进行比较研究。 本论文的数据来源于2004年5、8、11月和2005年2月在长江口北港和北支水域4个航次的海洋综合调查资料。通过采用优势度和优势种贡献率等生态学指标，同时结合逐步回归分析，讨论了温、盐与浮游动物总丰度变化的关系。结果表明，就四季而言，浮游动物总丰度和盐度呈显著正相关，和温度负相关。而就单个季节而言，除了5月和2月落潮，总丰度和盐度均呈显著正相关，与温度相关关系不显著。调查水域优势种主要为河口半咸水种或近海种。北港的盐度约为0-5, 表现为长江淡水特征；北支的盐度约为12-28，表现为长江径流和外海水团混合水团的特征。上述优势种的盐度适应和水团分布特征决定了长江口浮游动物分布的时空变化，即盐度较高的水域往往有较高的丰度，北支的总丰度和生物量均大于北港。此外，在长江口口内水域，水温较低的枯水期，浮游动物丰度高于水温较高的丰水期，温度的季节波动，以及主要优势种中华华哲水蚤（Sinocalanus sinensis）对低温环境的适应，是调查水域浮游动物总丰度季节变化的重要因素。 另一方面，本论文通过物种多样性、累积优势度曲线等群落水平的指标和相似性检验分析（ANOSIM），对长江口北港和北支浮游动物群落及其主要优势种的差异，以及主要生态因子对浮游动物群落的影响进行了分析。结果表明，北港主要由潮汐影响的淡水和低盐水组成，北支由中、高盐水组成。春季，北港浮游动物丰度和多样性均低于北支，主要优势种为中华华哲水蚤；北支优势种种数较多，主要是真刺唇角水蚤（Labidocera euchaeta）。夏季，北港和北支的多样性较接近，但是种类组成差别较大，北港以中华华哲水蚤和太平洋纺锤水蚤（Acartia pacifica）为主构成河口半咸水群落，北支以由火腿许水蚤（Schmackeria poplesia）、虫肢歪水蚤（Tortanus vermiculus）和太平洋纺锤水蚤等构成河口低盐群落。秋季，北港的丰度和多样性均低于北支，两个水域的群落差异明显，北港主要由中华华哲水蚤构成，北支由小拟哲水蚤（Paracalanus parvus）、针刺拟哲水蚤（Paracalanus aculeatus）、真刺唇角水蚤和中华哲水蚤（Calanus sinicus）等近岸和外海种组成。冬季，两个水域种数和多样性均较低，群落差别达四季最小，中华华哲水蚤占绝对优势。温度和盐度及其相互作用是决定长江口浮游动物群落特征的最主要因素。其中温度是造成研究水域浮游动物群落季节变化的主要因素，而由径流和外海水之间的消长引起的盐度变化是造成北港和北支浮游动物群落区域变化的决定因素。