The need for speed. I. Fast reactions and myelinated axons in copepods

A rapid and powerful escape response decreases predation risk in planktonic copepods. Calanoid copepods are sensitive to small and brief hydrodynamic disturbances: they respond with multiple nerve impulses to a vibrating sphere. Some species, such as Pleuromamma xiphias and Labidocera madurae, respond with very large spikes (1–4 mV), whereas maximum spike heights are an order of magnitude smaller in others, such as Undinula vulgaris and Neocalanus gracilis. A comparative study of the escape responses showed that all species reacted within 10 ms of the initiation of a hydrodynamic stimulus. However, U. vulgaris and N. gracilis had significantly shorter reaction times (minimum reaction times: 1.5 ms and 1.6 ms) than the other two, P. xiphias (6.6 ms) and L. madurae (3.1 ms). Examination of the first antenna and the central nervous system using transmission electron microscopy revealed extensive myelination of sensory and motor axons in the two species with the shorter reaction times. Axons of the other two species resembled typical crustacean unmyelinated fibers. A survey of 20 calanoids revealed that none of the species in two of the more ancient superfamilies possessed myelin, but myelination was present in the species from three more recently-evolved superfamilies. Accepted: 8 January 2000     

青鱼逃逸过程中的疾冲-滑行游泳行为

本实验以青鱼（Mylopharyngodon piceus）为研究对象,定性分析了通过电击实验鱼受到惊吓而产生的快速逃逸游泳行为,并定量分析了其疾冲游泳过程中加速度、最大疾冲游泳加速度和减速过程中的加速度。在自然光照下,保持实验水温为（20.0±1.0）℃。青鱼的3种不同体长规格分别为稚鱼（9.21±1.89）cm（n=30）、幼鱼（17.83±2.67）cm（n=30）和亚成体（61.45±0.80）cm（n=10）。实验过程中发现实验鱼逃逸游泳行为主要以疾冲-滑行的游泳方式进行,即实验鱼到达最大疾冲速度后身体保持固定直线不变的形式减速。3种实验体长的青鱼,其对应的最大绝对逃逸速度分别为（1.261±0.279）m/s、（1.542±0.280）m/s和（2.292±0.567）m/s,到达最大逃逸速度的时间分别为（0.249±0.089）s、（0.293±0.067）s和(0.216±0.024) s,对应的最大相对逃逸速度（实验鱼每秒内游泳距离相对于体长的倍数）分别为（13.694±3.032）BL/s、（8.648±1.571）BL/s和（3.729±0.923）BL/s。单因素方差分析表明,实验鱼的绝对疾冲速度随体长的增加而增加,亚成体的最大疾冲游泳速度显著大于稚鱼（P<0.05）;相对疾冲速度随体长的增加而减小,3种实验鱼之间的最大相对疾冲游泳速度均存在显著性差异（P<0.05）;亚成体的绝对滑行游泳加速度的绝对值显著高于幼鱼和稚鱼(P<0.05）。3种实验鱼之间的相对滑行加速度不存在显著性差异（P>0.05）。     

Adenosine and Prostaglandin E2 Cooperate in the Suppression of Immune Responses Mediated by Adaptive Regulatory T Cells

Adaptive regulatory T cells (Tr1) are induced in the periphery upon encountering cognate antigens. In cancer, their frequency is increased; however, Tr1-mediated suppression mechanisms are not yet defined. Here, we evaluate the simultaneous involvement of ectonucleotidases (CD39/CD73) and cyclooxygenase 2 (COX-2) in Tr1-mediated suppression. Human Tr1 cells were generated from peripheral blood mononuclear cell-derived, sorted CD4⁺CD25⁻ T cells and incubated with autologous immature dendritic cells, irradiated COX-2⁺ or COX-2⁻ tumor cells, and IL-2, IL-10, and IL-15 (each at 10–15 IU/ml) for 10 days as described (Bergmann, C., Strauss, L., Zeidler, R., Lang, S., and Whiteside, T. L. (2007) Cancer Immunol. Immunother. 56, 1429–1442). Tr1 were phenotyped by multicolor flow cytometry, and suppression of proliferating responder cells was assessed in carboxyfluorescein diacetate succinimidyl ester-based assays. ATP hydrolysis was measured using a luciferase detection assay, and levels of adenosine or prostaglandin E₂ (PGE₂) in cell supernatants were analyzed by mass spectrometry or ELISA, respectively. Intracellular cAMP levels were measured by enzyme immunoassay. The COX-2⁺ tumor induced a greater number of Tr1 than COX-2⁻ tumor (p < 0.05). Tr1 induced by COX-2⁺ tumor were more suppressive, hydrolyzed more exogenous ATP (p < 0.05), and produced higher levels of adenosine and PGE₂ (p < 0.05) than Tr1 induced by COX-2⁻ tumor. Inhibitors of ectonucleotidase activity, A_2A and EP₂ receptor antagonists, or an inhibitor of the PKA type I decreased Tr1-mediated suppression (p < 0.05), whereas rolipram, a PDE₄ inhibitor, increased the intracellular cAMP level in responder cells and their susceptibility to Tr1-mediated suppression. Tr1 present in tumors or the peripheral blood of head and neck squamous cell carcinoma patients co-expressed COX-2, CD39, and CD73. A concomitant inhibition of PGE₂ and adenosine via the common intracellular cAMP pathway might be a novel approach for improving results of immune therapies for cancer.     

Domestication reduces the capacity to escape from predators

Phenotypic plasticity in response to variations in predatory pressure frequently occurs in wild populations, but it may be more evident and critical in species subjected to high exploitation rates and aquaculture. The Chilean scallop Argopecten purpuratus is becoming a domesticated species and the production of hatchery-reared scallops (closed environment), has implied the development of successive generations of individuals deprived of several stimuli normally present in their natural habitats (e.g. predators). We compared the escape capacities between wild and cultured A. purpuratus and also evaluated the effect of reproductive investment on the escape response capacities. Wild and cultured scallops, at different reproductive stages (maturing, mature and spawned), were stimulated to escape with the predatory sea star Meyenaster gelatinosus. We recorded: (1) the time to reaction, (2) the total number of claps, the duration of the clapping response and the clapping rate until exhaustion, (3) the time they spent closed after exhaustion, and (4) the proportion of claps recovered, the duration of the clapping response and the clapping rate after 20 min of recuperation. We found that wild A. purpuratus (1) reacted earlier when contacted by their natural predator, (2) escaped faster (greater clapping rates), (3) spent less time with their valves closed when exhausted, and (4) most of their escape capacities (i.e. claps number; clapping time; capacity of recuperation) were less affected by the energetic requirements imposed by gonad maturation and/or spawning than in cultured scallops. We considered that all these aspects of the escape response would make wild scallops less vulnerable to predation than cultured scallops, thus decreasing predation risk. Given the reduction of escape performance in cultured scallops, we suggest that this aspect should be considered for the success of culture-based restocking programs.     

Complex interactions between a plant pathogen and insect parasitoid via the shared vector-host: consequences for host plant infection

Plant viruses modify the development of their aphid vectors by inducing physiological changes in the shared host plant. The performance of hymenopterous parasitoids exploiting these aphids can also be modified by the presence of the plant pathogen. We used laboratory and glasshouse microcosms containing beans (Vicia faba) as the host plant to examine the interactions between a plant virus (pea enation mosaic virus; PEMV) and a hymenopterous parasitoid (Aphidius ervi) that share the aphid vector/host Acyrthosiphon pisum. Neither PEMV-infection of V. faba, nor the carriage of PEMV virions by A. pisum, affected the growth or morphology of the aphid, or the oviposition behaviour and development of A. ervi. The presence of developing Aphidius ervi larvae within Acyrthosiphon pisum did not affect the ability of the aphids to transmit PEMV. However, by reducing their longevity, parasitism ultimately decreased the time viruliferous aphids were able to inoculate plants. In terms of virus dispersal, parasitized aphids exhibited more movement around experimental arenas than unparasitized controls, causing a slight increase in the proportion of beans infected with PEMV. Exposure to adult Aphidius ervi caused Acyrthosiphon pisum to rapidly drop off bean plants and disperse to new hosts, resulting in considerably higher plant infection rates (70%) than that seen in control arenas (25%). The results of this investigation demonstrate that when parasitoids are added to a plant-pathogen-vector system, benefits to the host plant due to reduced herbivore infestation must be balanced against the consequences of parasitoid-induced aphid dispersal and a subsequent increase in the level of plant infection.     

Fine-scale predation risk on elk after wolf reintroduction in Yellowstone National Park, USA

While patterns from trophic cascade studies have largely focused on density-mediated effects of predators on prey, there is increasing recognition that behaviorally mediated indirect effects of predators on prey can, at least in part, explain trophic cascade patterns. To determine if a relationship exists between predation risk perceived by elk (Cervus elaphus) while browsing and elk position within the landscape, we observed a total of 56 female elk during two summers and 29 female elk during one winter. At a fine spatial (0–187 m) and temporal scale (145–300 s), results from our model selection indicated summer vigilance levels were greater for females with calves than for females without calves, with vigilance levels greater for all females at closer escape-impediment distances. Winter results also suggested greater female vigilance levels at closer escape-impediment distances, but further indicated an increase in vigilance levels with closer conifer-edge distances. Placed within the context of other studies, the results were consistent with a behaviorally mediated trophic cascade and provide a potential mechanism to explain the variability in observed woody plant release from browsing in Yellowstone National Park, Wyoming, USA.     

Tonic contractions allow metabolic recuperation of the adductor muscle during escape responses of giant scallop Placopecten magellanicus

To escape from starfish predators, giant scallops, Placopecten magellanicus, swim using series of strong phasic contractions interrupted by tonic contractions. To investigate whether these tonic contractions allow metabolic recuperation of the adductor muscle, we sampled scallops at rest (Control), after an initial series of phasic contractions (Phasic) and after 1 min of tonic contraction following their initial phasic contractions (Phasic + Tonic) and compared muscle levels of phosphoarginine, adenylate nucleotides (ATP, ADP and AMP) and adenylate energy charge (AEC). Scallops in the two active groups did not differ in the numbers of phasic contractions or the mean phasic force production. Phosphoarginine concentrations in the adductor muscle decreased with phasic activity and remained low after 1 min of tonic contraction. ATP and ADP and total adenylate levels did not differ between the three groups, but AMP levels were higher in the scallops sampled after phasic contractions than in control scallops. The AEC was reduced by phasic contractions but returned to control levels after 1 min of tonic contraction. A significant negative correlation between AEC and the number of claps in the Phasic group disappeared in the Phasic + Tonic group. Thus, tonic contractions following phasic contractions allow partial metabolic recovery of the adductor muscle by returning AEC to control levels. However, phosphoarginine levels did not recover during tonic contractions, and a negative correlation between the number of claps and phosphoarginine levels remained in the Phasic + Tonic group. By interspersing tonic contractions between series of phasic contractions, scallops improved muscle energetic status, which should help maintain phasic force production during the remainder of the escape response.     

Force recordings during escape responses by Placopecten magellanicus (Gmelin): Seasonal changes in the impact of handling stress

Handling stress coupled with air exposure reduced the contractile performance during escape responses of 2+ scallops, Placopecten magellanicus, modifying maximal and mean phasic force production, the number of phasic contractions and the minimal interval between phasic contractions as well as reducing maximal tonic force production and increasing the reliance upon tonic contractions. The impact of handling stress was mitigated in scallops studied in the fall (late October) relative to the summer (late June and late August), with smaller declines in performance for all parameters examined. Seasonal changes in escape response performance by unstressed scallops were few, with virtually no change in maximal or mean phasic force production, in the number of phasic contractions or in the minimal interval between phasic contractions. As the scallops sampled in October had lower condition indices than those sampled in summer, the reduced impact of handling stress was likely due to the lower air and water temperatures. This suggests that transfers of juvenile scallops during culture operations should be done during cooler periods. Force measurements during escape responses are a sensitive, but simple tool with which to assess the status of scallops as they clearly reveal the marked impact of handling stress as well as more subtle seasonal changes.     

Escape reaction performance of myelinated and non-myelinated calanoid copepods

Calanoid copepods from seven families in three superfamilies were exposed to a controlled near-field hydrodynamic stimulus and their escape reactions were recorded using high-speed videographic techniques. Copepod species have two distinct mechanisms for increasing conduction speed of neural signals: larger diameter nerve axons and insulated axons, i.e., myelination. Myelinated axons have been found in certain species of the more recently-evolved calanoid superfamilies. Copepod representatives from these superfamilies were expected to have shorter response latencies than species from more ancestral superfamilies due to the increased conduction speed of nerve impulses in myelinated neurons. Using frame-by-frame playback and computerized motion analysis techniques, response latency, jump speed, and acceleration were measured. Kinetic performance of copepods was highly variable, with mean escape speeds ranging between 100-250 mm s^− 1 and accelerations of 9-230 m s^− 2. Minimum behavioral response latencies of 2 ms were recorded for both myelinated and non-myelinated calanoids. There was no significant difference between the response latencies of copepods from the myelinated and non-myelinated superfamilies. Furthermore, no relationships were found between copepod latency and size for either myelinated or non-myelinated species. Previous research may suggest that myelin may shorten the response latencies of certain calanoid species. However, our results show that non-myelinated copepods are also capable of responding rapidly, within as few as 2 ms, to hydrodynamic stimuli and produce similar kinetic performance to myelinated species. The main advantage of myelination over giant nerve axons is their more efficient transfer of nerve impulses resulting in a metabolic energy savings. Although this energetic reward would be important for copepods in food-limited environments, for coastal copepods, in food-rich habitats, either mechanism is a viable solution.