Experimental evaluation of herbivory in the ctenophore Mnemiopsis leidyi relevant to ctenophore-zooplankton-phytoplankton interactions in Narragansett Bay, Rhode Island, USA

Herbivory of Mnemiopsis leidyi and its interactions with phytoplanktonand non-gelatinous zooplankton were examined in small-scalemicrocosm experiments. Clearance rates for M. leidyi incubatedwith phytoplankton were generally negative, but ranged up to4.5 1 ctenophore^–1 day^–1 when the large (80 µmø) diatom Ditylum brightwelli was offered as a food source.These highest ingestion rates would provide Mnemiopsis withonly 21 % of its daily carbon requirements for respiration.Mean shrinkage of M. leidyi was 8.2–51% when incubatedwith phytoplankton. Although M. leidyi neither fed activelyon phytoplankton, nor satisfied its nutritional needs on sucha diet, the chain-forming diatom Skeletonema costatum becameentangled in mucus strands and balls produced by M. leidyi inthe absence of zooplankton. Attachment onto mucus occurred atphytoplankton concentrations commonly observed in NarragansettBay and may be important in the formation of "marine snow" duringsummer M. leidyi pulses; phytoplankton sinking rate and the"package size" available to herbivores would also be affected.The experiments support our previous hypothesis based on fieldobservations in Narragansett Bay that M. leidyi indirectly regulatesphytoplankton abundance there during the summer bloom as a consequenceof predation on zooplankton. The extent to which M. leidyi influencedphytoplankton dynamics in the microcosms was dependent on therelative abundance and physiological state of the three trophiclevels. A food web diagram for M. leidyi is presented.     

Life-stage dependent, in situ dietary patterns of the lobate ctenophore Mnemiopsis leidyi Agassiz 1865

Analysis of in situ gut contents of the ctenophore Mnemiopsisleidyi collected during summer months of 1998 at Woods Hole,MA, USA indicates that dietary changes accompanied developmentof this zooplankter. The transition from cydippid to lobatemorphologies involved a shift from a microplanktonic diet dominatedby protists to a metazoan-based diet. Dietary diversity increasedrapidly during the transition from cydippid to lobate stagebut remained essentially similar through most (>3.0 cm totallength) of the lobate stage. These stage-dependent dietary differencesmay have important implications for understanding populationdynamics of M. leidyi.     

Inductive interactions and embryonic equivalence groups in a basal metazoan, the ctenophore Mnemiopsis leidyi

Ctenophores undergo locomotion via the metachronal beating of eight longitudinally arrayed rows of comb plate cilia. These cilia are normally derived from two embryonic lineages, which include both daughters of the four e1 micromeres (e11 and e12) and a single daughter of the four m1 micromeres (the m12 micromeres). Although the e1 lineage is established autonomously, the m1 lineage requires an inductive interaction from the e1 lineage to contribute to comb plate formation. Successive removal of the e1 progeny at later stages of development indicates that this interaction takes place after the 32-cell stage and likely proceeds over a prolonged period of development. Normally, the e1, cell lies in closest proximity to the m12 cell that generates comb plate cilia; however, either of the e1 daughters (e11 or e12) is capable of emitting the signal required for m1 descendants to form comb plates. Previous cell lineage analyses indicate that the two e1 daughters generate the same suite of cell fates. On the other hand, the m1 daughters (m11 and m12) normally give rise to different cell fates. Reciprocal m1 daughter deletions show that in the absence of one daughter, the other cell can generate all the cell types normally formed by the missing cell. Together, these findings demonstrate that the two m1 daughters (m11 and m12) represent an embryonic equivalence group or field and that differences in the fates of the two m1 daughters are normally controlled by cell-cell interactions. These combined properties of ctenophore development, including the utilization of deterministic cleavage divisions, inductive interactions, and the establishment of embryonic fields or equivalence groups, are remarkably similar to those present in the development of various bilaterian metazoans.     

Regeneration of ciliary comb plates in the ctenophore Mnemiopsis leidyi. i. morphology

Regeneration of missing body parts in model organisms provides information on the mechanisms underlying the regeneration process. The aim here is to use ctenophores to investigate regeneration of their giant ciliary swimming plates. When part of a row of comb plates on Mnemiopsis is excised, the wound closes and heals, greatly increasing the distance between comb plates near the former cut edges. Video differential interference contrast (DIC) microscopy of the regeneration of new comb plates between widely separated plates shows localized widenings of the interplate ciliated groove (ICG) first, followed by growth of two opposing groups of comb plate cilia on either side. The split parts of a new plate elongate as their bases extend laterally away from the ICG widening and continue ciliogenesis at both ends. The split parts of a new plate grow longer and move closer together into the ICG widening until they merge into a single plate that interrupts the ICG in a normal manner. Video DIC snapshots of dissected gap preparations 1.5–3‐day postoperation show that ICG widenings and/or new plates do not all appear at the same time or with uniform spacing within a gap: the lengths and distances between young plates in a gap are quite variable. Video stereo microscopy of intact animals 3–4 days after the operation show that all the new plates that will form in a gap are present, fairly evenly spaced and similar in length, but smaller and closer together than normal. Normal development of comb plates in embryos and growing animals is compared to the pattern of comb plate regeneration in adults. Comb plate regeneration differs in the cydippid Pleurobrachia that lacks ICGs and has a firmer mesoglea than Mnemiopsis. This study provides a morphological foundation for histological, cellular, and molecular analysis of ciliary regeneration in ctenophores. J. Morphol. 2011. © 2011 Wiley Periodicals, Inc.     

Calcium-sensitive action potential of long duration in the fertilized egg of the ctenophore Mnemiopsis leidyi

The membrane properties of fertilized eggs of the ctenophore Mnemiopsis leidyi were studied using standard microelectrode techniques. The resting potential was approximately -80 mV, and was dependent on the extracellular K concentration. Depolarizing current injections elicited an action potential with an initial peak amplitude of +20 to +40 mV (duration about 5 sec) and a long lasting (duration 3 to 10 min) plateau phase. The depolarizing phase and the plateau phase appeared to have different ionic mechanisms. The entire action potential could be prevented by removal of extracellular Ca, but only the amplitude of the depolarizing phase, not the plateau phase, was dependent on the extracellular Ca concentration. The plateau phase was not observed in the absence of Ca, but in the presence of Ca its duration was dependent on the external Ca concentration. The data suggest that the plateau phase is activated as a consequence of Ca influx during the initial depolarizing phase. Removal of external Na resulted in only minor changes in the waveform of repolarization. The action potential was resistant to low concentrations of Mn and Cd in the presence of Ca. The role of this action potential in ctenophore development is not known, but in its waveform and duration it resembles the sperm-gated potentials that have been seen in eggs of other phyla. These experiments show ctenophore embryos to be excitable at very early stages, and suggest their utility in the study of the differentiation of cellular electrical properties.     

The ctenophore Mnemiopsis in native and exotic habitats: U.S. estuaries versus the Black Sea basin

The native habitats of the ctenophore, Mnemiopsis, are temperate to subtropical estuaries along the Atlantic coast of North and South America, where it is found in an extremely wide range of environmental conditions (winter low and summer high temperatures of 2 and 32 °C, respectively, and salinities of <2–38). In the early 1980s, it was accidentally introduced to the Black Sea, where it flourished and expanded into the Azov, Marmara, Mediterranean and Caspian Seas. We compile data showing that Mnemiopsis has high potentials of growth, reproduction and feeding that enable this species to be a predominant zooplanktivore in a wide variety of habitats; review the population distributions and dynamics of Mnemiopsis in U.S. waters and in the Black Sea region; and examine the effects of temperature and salinity, zooplankton availability and predator abundance on Mnemiopsis population size in both regions, and the effects of Mnemiopsis on zooplankton, ichthyoplankton and fish populations, focusing on Chesapeake Bay and the Black Sea. In both regions, Mnemiopsis populations are restricted by low winter temperatures (<2 °C). In native habitats, predators of Mnemiopsis often limit their populations, and zooplanktivorous fish are abundant and may compete with the ctenophores for food. By contrast, in the Black Sea region, no obvious predators of Mnemiopsis were present during the decade following introduction when the ctenophore populations flourished. Additionally, zooplanktivorous fish populations had been severely reduced by over fishing prior to the ctenophore outbreak. Thus, small populations of potential predators and competitors for food enabled Mnemiopsis populations to swell in the new habitats. In Chesapeake Bay, Mnemiopsis consumes substantial proportions of zooplankton daily, but may only noticeably reduce zooplankton populations when predators of Mnemiopsis are uncommon. Mnemiopsis also is an important predator of fish eggs in both locations. In the Black Sea, reductions in zooplankton, ichthyoplankton and zooplanktivorous fish populations have been attributed to Mnemiopsis. We conclude that the enormous impact of Mnemiopsis on the Black Sea ecosystem occurred because of the shortage of predators and competitors in the late 1980s and early 1990s. The appearance of the ctenophore, Beroe ovata, may promote the recovery of the Black Sea ecosystem from the effects of the Mnemiopsis invasion.     

青岛崂山青山湾人工鱼礁区及附近海域鱼卵仔稚鱼种类组成与数量分布

为研究人工鱼礁对产卵鱼类的诱集效果和庇护效应,2014—2015年对青岛崂山青山湾海域人工鱼礁区及附近海域鱼卵和仔稚鱼的种类组成和数量分布进行了水平和垂直拖网调查. 2014年春、夏、秋3个季节的7个航次调查中,共采集鱼卵7306粒、仔稚鱼52尾,隶属于4目9科11属12种;2015年同期进行的13个航次调查中,共采集鱼卵10373粒、仔稚鱼159尾,隶属于6目11科14属15种.2年间鱼卵和仔稚鱼样品均以鲈形目数量最多,鲽形目次之,其中鱼卵样品中仅有少鳞鱚和短吻红舌鳎构成优势种;仔稚鱼样品以少鳞鱚优势度最高,但不存在仔稚鱼优势种.鱼卵和稚鱼的高优势度物种的季节间演替明显.春、夏、秋季鱼类浮游生物群落Margalef丰富度指数、Shannon多样性指数和Pielou均匀度指数平均值均较低,说明该海域鱼类浮游生物群落结构稳定性较低.2年间所采集物种对应的成鱼平均营养级分别为3.71和3.78,均属第三营养级中的低级肉食性鱼类,绝大多数为暖温种,无冷温性种类,表现出明显的暖温带特点.综合分析认为,人工鱼礁区鱼类浮游生物群落的多样性相对较高,这与鱼礁区流速和流场特征及复杂的空间异质性有关.     

Growth and feeding rates of the newly hatched larval ctenophore Mnemiopsis leidyi A. Agassiz (Ctenophora, Lobata)

Mnemiopsis leidyi: larvae depend on microplankton (<200 µm) prey duringthe first few days following hatching until larvae are >0.5mm in length and can successfully capture and consume mesozooplanktonprey. Feeding and growth rates of newly hatched M. leidyi larvaewere measured in controlled laboratory experiments. When fednatural microplankton assemblages, newly hatched larvae consumedsignificant quantities of both autotrophic and heterotrophicprey, including diatoms, phototrophic, heterotrophic and mixotrophicdinoflagellates, euglenoid flagellates, aloricate and tintinnidciliates, and rotifers. Average per capita clearance rates were1.99–7.59 mL individual^–1 h^–1 ( = 4.01 mL individual^–1 h^–1; SD = 1.95)and total per capita ingestion was 0.01–4.70 µgC individual^–1 day^–1 x 10² ( = 0.83 µg C individual^–1 day^–1 x 10²; SD =1.89). Larval growth rates were –0.13 to 0.56 mm individual^–1day^–1 (equivalent to –1.72 to 4.33 µg C individual^–1day^–1) over a range of larval sizes from 0.5 (<0.5µg C) to 5 mm (85 µg C). A diet consisting entirelyof microplankton prey supported larval growth for >2 weeks,and growth rate decreased when larvae reached 4–5 mm inlength, corresponding to the beginning of their morphologicaltransition from tentaculate to lobate feeding mode. The grossgrowth efficiency of larvae fed natural microplankton assemblageswas 3%.     

Distribution of ichthyoplankton in the eastern Peter the Great Bay,Sea of Japan,in June and July of 2007

Comparison of the results of ichthyoplankton surveys conducted at 97 stations in the eastern part of the Peter the Great Bay of the Sea of Japan, in June–July 2007 with the similar research data of the 1950s shows that at present, as was the case 50 years ago, flatfish eggs belonging mainly to the yellowfin sole Limanda aspera and brown sole Pleuronectes herzensteini prevail in the local ichthyoplankton (up to 86%). The highest concentrations of these species’ eggs were recorded in the Vostok Bay and Strelok Bay. The spawning activity of flatfish in 2007 is found to be lower than in the mid 1900s, but the significance of the eastern part of the Peter the Great Bay for flatfish reproduction remains large. The importance of long-term monitoring in this area, which is being subjected to steadily growing anthropogenic impacts, is also proven.     

Vulnerability of late larval and early juvenile Atlantic herring, Clupea harengus, to predation by whiting,Merlangius merlangus

Laboratory experiments investigated changes in the vulnerability of herring to predation by whiting during growth from <34 to >80mm in length, Premetamorphic herring (<50mm) failed to react to c . 50% of attacks by the predator, but this was reduced to c . 20% in postmetamorphic fish. Premetamorphic herring failing to react had c. 30% probability of survival due to unforced errors by the predator (unsuccessful attacks that did not elicit a reaction by the prey), but these errors did not occur after herring metamorphosis. The loss of the advantage of lower conspicuousness following metamorphosis was counterbalanced by increased reactivity. Antipredator benefits should increase with schooling behaviour (first evident in experiments at c . 50 mm), so predation mortality should decrease in postmetamorphic herring in the sea. Metamorphosis itself would be associated with high predation risk since conspicuousness is enhanced but reactivity and schooling behaviour are still not fully developed.     

Laboratory studies of predation by the ctenophore Mnemiopsis leidyi on the early stages in the life history of the bay anchovy, Anchoa mitchilli

The lobate ctenophore, Mnemiopsis leidyi, consumed eggs andlarvae of the bay anchovy, Anchoa mitchilli, in laboratory experiments.This ctenophore exhibited a type I functional response to increasesin egg densities without reaching saturation at high prey densities.Clearance rate increased with increasing experimental containervolume. There was a 3-fold increase in the volume of water clearedby 2.0–2.5-cm ctenophores and a >5-fold increase for4.5–5.0-cm ctenophores in 15–1 versus 100–200–1containers. Clearance rate was dependent on tenophore lengthbut was probably underestimated for the larger animals due tocontainer effects. The presence of various densities of alternateprey, Acartia hudsonica or Anemia sp. nauplii, in addition toAnchoa mitchilli eggs did not affect the clearance rates onthe eggs alone. Comparison of clearance rates of 2.0–2.5-cmctenophores on various ages of starved and fed bay anchovy larvaeindicated that predation may be higher on yolk-sac larvae thanon eggs but decreases as the larvae grow. After 3 days posthatch starved anchovy larvae become more vulnerable to predationthan fed larvae. The ctenophore, M.leidyi has the potentialto inflict substantial predation pressure on early stages inthe life history of bay anchovy. ¹ Present address: University of Maryland, Center for Environmentaland Estuarine Studies, Chesapeake Biological Laboratory, POBox 38, Solomons, MD 20688-0038, USA     

Seasonal changes and population dynamics of the ctenophore Mnemiopsis leidyi after its first year of invasion in the Kiel Fjord, Western Baltic Sea

We analyzed the seasonal variations of the ctenophore Mnemiopsis leidyi weekly collected since its first record in the western Baltic Sea in October 2006. The distribution pattern together with the seasonal dynamics and population outbreaks in late summer 2007 indicate recent successfully establishment of M. leidyi in this area. Seasonal changes showed two periods of high reproductive activity characterized by a population structure dominated by small size classes, followed by an increase of larger ones. These results further revealed that the bulk of the population remains in deep layers during the periods of low population density, whereas it appeared situated in upper layers during the proliferation of the species. We further emphasized the strength of the population outbreaks, which can reach abundances >10-fold higher in time periods shorter than a week. The predatory impact this species may have in pelagic ecosystems warns on the importance of its recent range of expansion.     

Intracellular fate mapping in a basal metazoan, the ctenophore Mnemiopsis leidyi, reveals the origins of mesoderm and the existence of indeterminate cell lineages.

Ctenophores are marine invertebrates that develop rapidly and directly into juvenile adults. They are likely to be the simplest metazoans possessing definitive muscle cells and are possibly the sister group to the Bilateria. All ctenophore embryos display a highly stereotyped, phylum-specific pattern of development in which every cell can be identified by its lineage history. We generated a cell lineage fate map for Mnemiopsis leidyi by injecting fluorescent lineage tracers into individual blastomeres up through the 60-cell stage. The adult ctenophore body plan is composed of four nearly identical quadrants organized along the oral-aboral axis. Each of the four quadrants is derived largely from one cell of the four-cell-stage embryo. At the eight-cell stage each quadrant contains a single E ("end") and M ("middle") blastomere. Subsequently, micromeres are formed first at the aboral pole and later at the oral pole. The ctene rows, apical organ, and tentacle apparatus are complex structures that are generated by both E and M blastomere lineages from all four quadrants. All muscle cells are derived from micromeres born at the oral pole of endomesodermal precursors (2M and 3E macromeres). While the development of the four quadrants is similar, diagonally opposed quadrants share more similarities than adjacent quadrants. Adult ctenophores possess two diagonally opposed endodermal anal canals that open at the base of the apical organ. These two structures are derived from the two diagonally opposed 2M/ macromeres. The two opposing 2M/ macromeres generated a unique set of circumpharyngeal muscle cells, but do not contribute to the anal canals. No other lineages displayed such diagonal asymmetries. Clones from each blastomere yielded regular, but not completely invariant patterns of descendents. Ectodermal descendents normally, but not always, remained within their corresponding quadrants. On the other hand, endodermal and mesodermal progeny dispersed throughout the body. The variability in the exact complements of adult structures, along with previously published cell deletion experiments, demonstrates that cell interactions are required for normal cell fate determination. Ctenophore embryos, like those of many bilaterian phyla (e.g., spiralians, nematodes, and echinoids), display a highly stereotyped cleavage program in which some, but not all, blastomeres are determined at the time of their birth. The results suggest that mesodermal tissues originally evolved from endoderm tissue.     

Large but uneven reduction in fish size across species in relation to changing sea temperatures

Ectotherms often attain smaller body sizes when they develop at higher temperatures. This phenomenon, known as the temperature–size rule, has important consequences for global fisheries, whereby ocean warming is predicted to result in smaller fish and reduced biomass. However, the generality of this phenomenon and the mechanisms that drive it in natural populations remain unresolved. In this study, we document the maximal size of 74 fish species along a steep temperature gradient in the Mediterranean Sea and find strong support for the temperature–size rule. Importantly, we additionally find that size reduction in active fish species is dramatically larger than for more sedentary species. As the temperature dependence of oxygen consumption depends on activity levels, these findings are consistent with the hypothesis that oxygen is a limiting factor shaping the temperature–size rule in fishes. These results suggest that ocean warming will result in a sharp, but uneven, reduction in fish size that will cause major shifts in size‐dependent interactions. Moreover, warming will have major implications for fisheries as the main species targeted for harvesting will show the most substantial declines in biomass.     

Timing of diapause termination in relation to variation in winter climate

In temperate insects, winters are typically endured by entering diapause, which comprises a deep resting stage. Correct timing of diapause termination is vital for synchronization of emergence with conspecifics and for mobilizing resources when conditions for growth and reproduction become favourable. Although critical to survival, the intrinsic and extrinsic drivers of diapause termination timing are poorly understood. In the present study, we investigate diapause development under a range of durations (10–24 weeks) spent at different temperatures (?2 to 10 °C) in the pupal diapausing butterfly Pieris napi Linnaeus (Lepidoptera:Pieridae). We determine: (i) the maximum cold temperature for diapause development; (ii) if pupae in diapause count cold days or cold sums; and (iii) whether diapause termination is distinct or gradual. The results indicate large and idiosyncratic effects of high and low nonlethal temperatures on diapause development in P. napi. Although all temperatures tested lead to diapause termination, a thermal optimum between 2 and 4 °C is observed. Lower temperatures lead to decreased eclosion propensity, whereas higher temperatures slow down development and increase emergence desynchronization. These data suggest that, rather than a simple cold‐summing process with a distinct diapause termination point, there are trade‐offs between time and temperature at the low and high end of the thermal range, resulting in a nonlinear thermal landscape showing a ridge of increasing eclosion propensity at moderate temperatures. The present study suggests that the effects of temperature on diapause development should be included in projections on post‐winter phenology models of insects, including pest species.     

长江口水域夏季鱼卵和仔稚鱼年间变化

基于2005年、2008年、2009年和2011年8月(夏季)在长江口水域(30°30'—31°45'N,121°15'—123°10'E)4个航次的浮游生物拖网资料,分析了长江口水域鱼卵和仔稚鱼的种类组成、数量分布特征及其年间变化。结果表明:4个航次采集的鱼卵和仔稚鱼鉴定到种的种类有17种,隶属于8目13科,以鲈形目种类最多,11种,其次是鲱形目,5种,其他各目种类均小于5种;种类数存在明显年间差异,2005年种类数最多(鱼卵3种,仔稚鱼8种),其次是2009年和2011年,2008年种类数最少(鱼卵1种,仔稚鱼5种)。优势种年间更替明显,长蛇鲻(Saurida elongata)、虾虎鱼(Gobiidae spp.)和中华小公鱼(Stolephorus chinensis)在2005年是优势种,2008年优势种为鳀鱼(Engraulis japonicus),2009年优势种为鳀鱼、寡鳞飘鱼(Pseudolaubuca engraulis)、虾虎鱼等,2011年虾虎鱼和小公鱼(包括小公鱼属未定种Stolephorus spp.和中华小公鱼Stolephorus chinensis)成为优势种。2005年鱼卵和仔稚鱼数量分布的密集区在嵊泗列岛附近水域,2008年鱼卵和仔稚鱼出现较少,未出现明显的数量密集区;2009年鱼卵数量较少,仔稚鱼数量较多,密集区主要分布在在长江口以外123°E附近水域;2011年鱼卵主要分布在在长江北支口门外附近水域,仔稚鱼在调查区内分布相对均匀。