Methods of ciliary filter feeding in adult Phoronis muelleri (phylum Phoronida) and in its free-swimming actinotroch larva

Microscope-video recordings were used to describe basic characteristics of particle capture in adult Phoronis muelleri and in its actinotroch larva. The larva captures food particles by means of close interplay between the tentacles and the oral hood. Suspended particles in the incoming water flow set up by the lateral "pump" cilia on the tentacles approach the tentacles with a velocity of about 1.5 mm s^-1. Near the tentacles, the particles are stopped by the stiff sensory laterofrontal cilia acting as a mechanical sieve. Simultaneous with stoppage of a particle, the oral hood is rapidly lifted so that water from the surrounding area flows in to occupy the volume created under the hood. Due to the proximity of the tentacle with the arrested particle, this suction will draw the particle away from the laterofrontal cilia into the increased space under the oral hood with a velocity of about 0.6 mm s^-1 before it is subsequently carried to the mouth. If a particle is stopped near the tip of a tentacle this may trigger a tentacle flick, with a tip velocity of about 5-7 mm s^-1, which brings the particle down toward the lifting edge of the hood. A stopped particle may cause a local disruption of the metachronism of the lateral cilia for about 0.14 s. Likewise in the adult, when an incoming particle with a velocity of about 1.6 mm s^-1 is stopped near the tip of a tentacle this triggers a flick, which brings the particle down towards the mouth. The duration of the active flick phase is about one-tenth of the flick cycle. Only when a particle is stopped on the outer part of a tentacle is a flick triggered. Otherwise the particle is either transported down along the frontal surface of the tentacle by means of the frontal cilia, or transferred into the downward current created by the lophophore. The metachronal wave velocity is about 0.25 mm s^-1, the wavelength about 12 µm, and hence the ciliary beat frequency about 21 Hz (at ~16°C). Essential features of the filter-feeding process in both actinotroch larvae and adult P. muelleri reported here resemble the ciliary-sieving process described for bryozoans in recent years. It should be pointed out that the stiff, mechanosensory laterofrontal cilia may be a decisive factor for understanding ciliary upstream collecting, also in pterobranchs and brachiopods in which sensory laterofrontal cilia are present.     

Energetic costs of swarming behavior for the copepod Dioithona oculata

The cyclopoid copepod Dioithona oculata forms dense swarms within shafts of sunlight that penetrate the mangrove prop-root habitat of islands off the coast of Belize. Previous studies, based on in situ video recordings and laboratory studies, have shown that D. oculata is capable of maintaining fixed-position swarms in spite of currents of up to 2 cm s⁻¹. The purpose of this study was to examine the energetic costs of maintaining these swarms, in terms of increased metabolic costs of maintaining position in currents and in terms of reduced feeding rates in densely packed swarms during the day. Using a sealed, variable-speed flow-through chamber, the respiration rates of D. oculata were measured while swarms maintained position in different current speeds. The results indicate that active metabolism (swimming at maximum speed to maintain the swarm in a current) is approximately three times greater than routine metabolism (normal swimming speeds in the absence of currents), indicating a significant metabolic cost of maintaining swarms in the presence of currents. In addition, gut-pigment analysis indicated that feeding rates of these copepods were often reduced in swarms during the day compared to when the copepods were dispersed at night. Given the high “cost” of swarming, the adaptive value of swarming in terms of reduced predation, increased opportunities for mating, and reduced dispersal, must be substantial. Received: 4 June 1997 / Accepted: 18 September 1997     

Energetic and time costs of foraging in harvester ants,Pogonomyrmex occidentalis

Summary Western harvester ants, Pogonomyrmex occidentalis, preferentially utilize low vegetational cover pathways. Energetic costs for foraging ants were less than 0.1% of caloric rewards of harvested seeds, suggesting that reduction of energetic cost is not a major benefit of this preference. Walking speed was significantly faster on lower cover routes, increasing net return rates from equidistant artificial food sources. Undisturbed foragers on low cover routes traveled farther, increasing their total foraging area without increasing foraging time. These results suggest that in animals with low costs of locomotion relative to energetic rewards, time costs are more important than direct energetic costs in influencing foraging decisions. In baited experiments with equidistant food sources, preferential use of low cover routes resulted in a large increase in net energetic gain rate, but only a slight increase in energetic efficiency. Under natural conditions, net energetic gain rates were constant for foragers using low and high vegetational cover routes, but foragers using low cover paths had lower efficiencies. This suggests that net energetic gain rate is a more important currency than energetic efficiency for foraging harvester ants.     

Nuptial feeding in tettigoniids male costs and the rates of fecundity increase

Summary A recent debate has centred on the importance of paternal investment for the origin and maintenance of nuptial feeding in insects. Some authors have argued that the rates of nutrient incorporation are likely to be too slow to allow a male to fertilize the eggs that he helps to produce and cannot be considered as paternal investment. Here I report the results of some experiments that show that the positive effects of nutrient donation on female reproduction for one species of spermatophylax producing tettigoniid can be realized within 24 to 48 h. Furthermore, mating was found to induce a refractory period in the female that was longer than the time taken for the incorporation of nutrients and oviposition. Thus, the nurturant male is likely to fertilize the eggs even if last-male sperm precedence is high. The cost of spermatophylax production was manifest as a 5-day recovery period between matings. Thus males and their donations are likely to be limited resources for females resulting in a reversal of the typical sex roles.     

Particle capture in the crown of the ciliary suspension feeding polychaete Sabella penicillus: videotape recordings and interpretations

An experimental setup was designed for in situ videotape recording of the particle capture process in the crown of the polychaete Sabella penicillus. Intact individuals of S. penicillus (collected in the Gullmornfjord, Sweden in 1992) were exposed to either 6 m Latex spheres or Rhodomonas sp. flagellate cells (6 m). The capture of the added particles was recorded on video. From frameby-frame analyses particle velocities were estimated and the shape of the three-dimensional particle paths was inferred. The mean velocity of particles approaching the crown was estimated to be ca. 1 mm s^-1, increasing to ca. 1.7 mm s^-1 in the interpinnule channel. At the moment of capture the particles were seen to follow a curved, near circular path close to the tips of the latero-frontal cilia. The transport velocities on the frontal side of the pinnules and filaments were estimated to be up to 0.15 and 0.5 mm s^-1, respectively. Counting captured particles relative to particles arriving within the area of the pinnules gave a rough, direct estimate of nearly 100% retention rate when the polychaete was feeding undisturbed. Together with results from clearance measurements in the literature this implies that the worm is able to capture particles down to 3 m entering the interpinnule channel almost 100% effectively. In view of the 80-m wide interpinnule channel and 40-m spacing between the tips of the latero-frontal cilia on both sides of the channel, this result cannot be explained by mechanisms based solely on direct mechanical contact between cilia and particles but must involve fluid mechanical mechanisms. The present work is the experimental basis for ongoing numerical simulations of the particle motion in the interpinnule channel.     

Digestion rates and assimilation efficiencies of siphonophores fed zooplankton prey

Digestion times and assimilation efficiencies are critical factors used in calculations of carbon and nitrogen budgets. Digestion times of natural copepod prey differed significantly among 4 genera of siphonophores (P>0.001), from a minimum of 1.6 h to a maximum of 9.6 h. Assimilation efficiencies, in contrast, were uniformly high; 87 to 94% for carbon and 90 to 96% for nitrogen. Nitrogen assimilation was consistently greater than carbon assimilation. Assimilation efficiencies calculated according to dry weight substantially underestimated assimilation of carbon and nitrogen, while calculations using ash-free dry weights and ash-free to dry weight ratios approached values for carbon assimilation. These values are appreciably higher than most of the assimilation efficiencies previously measured for a few other planktonic carnivores. These results indicate very efficient digestion of food by siphonophores in oceanic environments where prey densities are low.     

Reanalysis of conversion efficiencies for larval Fundulus heteroclitus

The conversion efficiency (gross growth efficiency) of larval Fundulus heteroclitus fed newly hatched Artemia sp. nauplii was recently reported as the lowest observed (0.69 to 1.11%) for a fish. A recalculation of the efficiencies, based on data from Radtke and Dean, showed that the original determinations were in error by two orders of magnitude. The recalculated values (69.0 to 114.4%), however, are impossibly large and probably result from measurement error in the determination of the weight of a single prey individual (nauplius). Using a nauplius weight estimate from the literature, the conversion efficiencies were again recalculated. The new efficiencies (25 to 41%) agree with those reported for other fish species. The values are still higher than those reported elsewhere for adult F. heteroclitus, but this is consistent with the generalization that conversion efficiency decreases with age.Contribution No. 422 of the University of Georgia Marine Institute     

Some costs of reproduction for male bushcrickets,Requena verticalis (Orthoptera : Tettigoniidae) allocating resources to mate attraction and nuptial feeding

Summary The cost of reproductive effort is known to result in a trade-off between current and future reproduction. Similarly, trade-offs in energy allocation may occur between components of reproductive effort, mating and parental effort, within a single reproductive episode. We investigated the energy allocated to mating effort (calling to attract females) and parental effort (donation of spermatophore nutrients at mating) by male bushcrickets, Requena verticalis, under two dietary regimes. Males provided with a low quality diet reduced the daily energy allocated to calling activity while maintaining their investment in spermatophores. Males provided with a high quality diet did not allocate more resources per day to their spermatophores but stored excess energy for future reproduction. Thus, on a per day basis, males appear to hold constant their investment in the spermatophore at the cost of reduced mating effort when resources are limited. Males on both diets, however, increased the size of their spermatophore donations when the interval between female encounters was increased. One explanation for this pattern could be a frequency-dependent optimization of spermatophore size. Correspondence to: L.W. Simmons     

Time and recruitment costs as currencies in manipulation studies on the costs of reproduction

Life history theory predicts that parents will have lower Darwinian fitness if they tend clutches that are above or below the size they naturally produce. We experimentally tested for relationships between fitness and clutch size in Tree Swallow (Tachycineta bicolor) offspring and parents. Over 130 trios of nests initiated on the same day were randomly divided among reduce (-3 eggs), control (3 eggs picked up and replaced), or add (+3 eggs) manipulations. Pre-manipulation modal clutch size was six eggs (range before manipulations was 1-10; afterwards, it was 1-11). Hatching took longer in larger clutches, but the proportion of eggs hatching and fledging was similar for clutches from 4 to 10, so that clutches of 10 produced the maximum number of fledgling. Parental feeding rates were higher for larger broods, but per capita feeds to nestlings were fewer, and nestlings were smaller. Nonetheless, survival of both young and adults, based on recaptures in subsequent years, was not significantly affected by manipulations. Manipulations also had no significant effect on subsequent reproduction, including the number of fledglings produced by either local recruits or returning breeders. Collectively, our results failed to detect fitness costs associated with tending larger clutches for either parents or the offspring reared and suggested directional selection for larger clutch size. However, because clutches that hatch later produce fewer recruits, the extra days required to lay more eggs and to fledge extra young may eliminate a large part of the advantage that would accrue to parents producing enlarged clutches. For example, our data suggest that there may be less than a 16% benefit to producing nine instead of six eggs, rather than 50%, as is suggested by experimentally manipulated egg numbers alone. Thus, time, rather than costs of reproduction, may be the crucial constraint selecting against Tree Swallows laying larger clutches.     

复合二氧化氯对药物及个人护理品的降解效能

药物及个人护理品(PPCPs)在水处理工艺中的迁移转化备受关注.二氧化氯(ClO_2)是一种常用的消毒剂,但水厂中二氧化氯制备工艺得到的常为复合二氧化氯(即ClO_2和Cl_2的混合物).目前,复合二氧化氯对PPCPs的去除方面研究还比较缺乏.本文研究了复合二氧化氯对16种常见PPCPs的降解效能,并与单独二氧化氯氧化进行比较.研究发现,PPCPs的降解效果与其化学结构密切相关.ClO_2可选择性氧化磺胺甲基嘧啶等含富电子基团的PPCPs,对甲硝唑等不含富电子基团的PPCPs降解能力有限,对咖啡因和扑米酮基本不降解.相比单独二氧化氯氧化,复合二氧化氯能促进萘啶酸、氟甲喹和对乙酰氨基酚的降解,但是对甲硝唑、二甲硝咪唑和避蚊胺的降解能力低于单独二氧化氯氧化.复合二氧化氯使用中亚氯酸盐生成量降低.     

Feeding,conversion efficiencies,and growth of larval mummichogs,Fundulus heteroclitus

Feeding rates, conversion efficiencies and growth of larvae of the mummichog Fundulus heteroclitus, an extremely abundant estuarine fish, were measured at temperatures ranging from 18° to 30°C. The food used was Artemia salina nauplii. At the time of total yolk sac absorption (5 to 7 days after hatching), the feeding rate decreased for a short time, an indication of a shift in metabolism. Higher feeding rates and growth occurred at higher rearing temperatures. The highest conversion efficiency (gross growth efficiency) was 1.1%, at 22°C. Mummichog larvae may be energetically inefficient compared with other fish species, but efficiency might not be critical for this fish, which is an opportunistic omnivore in an energy-rich environment.Contribution No. 291 of the Belle W. Baruch Institute for Marine Biology and Coastal Research, supported by DOE contract No. EY-76-5-09-0869.     

A comparison of absorption and assimilation efficiencies between four species of shallow- and deep-living fishes

We captured two species of deep-sea zoarcids, Melanostigma pammelas and Lycodapus mandibularis, from Monterey Bay California and maintained them in the laboratory. One shallow-water zoarcid, Eucryphycus californicus, and an ecologically and morphologically similar stichaeid fish Xiphister atropurpureus were collected from intertidal and subtidal habitats in Monterey Bay. We investigated the absorption and assimilation efficiencies of these fishes to determine whether deep-sea species have evolved mechanisms to increase their efficiency of food use. Fishes were placed in experimental chambers and fed a known quantity of food. Ammonia excretion was measured and feces were collected daily. Both food and feces were analyzed for water, protein, lipid and ash to determine specific absorption efficiencies. Absorption ranged from 87.7 to 92.4% and assimilation efficiencies from 84.0 to 86.5%. Meal sizes from 0.5 to 4.0% of body weight did not affect the results. No significant differences were found between deep-sea and shallow-water species fed single meals or fed ad libitum for 10 days. This suggests that the selective pressure to maximize absorption and assimilation is universal and is not affected by the productivity of the habitat occupied. However, the relative size of the intestine in the deep-sea species was significantly smaller suggesting that they had a lower metabolic cost to maintain their digestive apparatus. It could not be concluded whether this was the result of pressure to conserve energy or rather the tendency of the shallow-living species to ingest more refractory material (i.e. sediment, algae).     

Ecological success of the cladoceran <Emphasis Type="Italic">Penilia avirostris</Emphasis> in the marine environment: feeding performance,gross growth efficiencies and life history

Marine cladocerans are important contributors to the zooplankton community of tropical and temperate coastal ecosystems during the warmer months, when they show explosive population growth. Despite this fact, little information is available on their ecology compared with the extensive studies on their freshwater relatives. The main objective of this study was to determine the in situ feeding and growth rates, and life history parameters of Penilia avirostris in São Sebastião Bay (Brazil) during austral summer 2004, as a premise to understand the advantages of this cladoceran in oligotrophic waters. Culture development experiments, monitored for a period of 12 days, showed that maximum juvenile release occurred after 2 days, and that the development duration of a complete cohort was around 9 days. From bottle incubation grazing experiments, significant ingestion rates upon flagellates, ciliates, dinoflagellates and diatoms were detected. Flagellates were the most important contributors to P. avirostris diet (ca. 80%). P. avirostris ingested between 28 and 97% of its own carbon biomass per day (daily ration) and individual growth rates of this marine cladoceran (0.10–0.24 d^?1) increased with prey availability. The combination of ingestion rates of natural prey and growth rates provided gross growth efficiencies (GGE) of 15–53%, on a carbon basis. Our results suggest that P. avirostris has similar GGE to copepods, although at low food conditions the values for the marine cladocerans seems slightly higher. However, this characteristic alone does not explain the explosive growth and community dominance shown by P. avirostris. Therefore, other traits related to the reproductive biology of the species, such as short generation time, parthenogenetic reproduction, and continuous somatic growth, seems to be mostly responsible for the success of P. avirostris in many marine ecosystems during their seasonal occurrence.     

Adjustment costs and optimal waste treatment

Consideration is given to various reasons for changing standards on emissions, effluents, and ambient air and water quality. It is proposed that adjustment costs may be of significant practical importance in determining the optimal time path of waste treatment (at least currently). The problem of minimizing the present value of the sum of treatment plus damage plus adjustment costs is placed in a calculus of variations framework and the optimal time path of waste treatment is characterized as well as the corresponding optimal tax. An example is used to illustrate some features of the solution.     

Zooplankton feeding and queueing theory

Three of the most commonly used mathematical relationships between zooplankton ingestion and food concentration are derived from a queue model, by assuming different feeding strategies of the individual predators or grazers. The model suggests that the abruptness of the transition between maximum search or filtration rate and maximum ingestion rate depends on the size ratio between the animals and their food items. The results show that the stochastic nature of predation and grazing cannot be neglected when the food particles are relatively big.     

Global justice and the opportunity costs of conservation

Opportunity costs can represent a significant portion of the costs associated with conservation projects and frequently outstrip other kinds of cost. They are typically understood to refer to the benefits someone would have obtained if conservation projects had not required them to give up current activities, such as farming or hunting or if the land had been available for uses other than conservation. This familiar way of identifying opportunity costs is flawed, however, because it threatens to condone, or take advantage of, the injustices that many people face that affect their opportunities. I integrated ideas from the political theory of global justice to examine how the analysis of opportunity costs illustrates the importance of considering conservation and issues of global justice together, rather than thinking about them in isolation. I distinguish four baselines for defining opportunity costs. A status quo baseline defines opportunity costs by asking what people would have earned had a conservation project not happened. A willingness to accept baseline defines them by asking people what it would take to make them indifferent to whether a conservation project takes place or not. An antipoverty baseline suggests that opportunity costs have been met when people affected by a project are not left in poverty. An egalitarian baseline suggests opportunity costs have been met when people are not left in relative disadvantage, with worse than average opportunities. I argue that the egalitarian baseline is the most acceptable from the point of view of justice. Such a baseline would suggest that, in practice, many of the world's poor are being unjustly treated, or even exploited, as a result of conservation activities.     

Resource extraction and recycling with environmental costs

Extraction and use of a natural resource is assumed to affect the environment adversely. A perfect substitute for the resource can be supplied through a recycling process. Recycling may also have harmful effects on the environment, but to a smaller extent than extraction. The optimal path of extraction and recycling is studied under various assumptions about the environmental effects of recycling and the assimilative capacity of the environment. In particular, it is shown how the cost of recycling will affect initial resource extraction as well as the environmental quality at the time of resource exhaustion and in the long-run stationary state.     

On costs and decisions in population management

Many populations have a negative impact on their habitat, or upon other species in the environment, if their numbers become too large. For this reason they are often managed using some form of control. The objective is to keep numbers at a sustainable level, while ensuring survival of the population. Here we present models that allow population management programs to be assessed. Two common control regimes will be considered: reduction and suppression. Under the suppression regime the population is maintained close to a particular threshold through near continuous control, while under the reduction regime, control begins once the population reaches a certain threshold and continues until it falls below a lower pre-defined level. We discuss how to best choose the control parameters, and we provide tools that allow population managers to select reduction levels and control rates. Additional tools will be provided to assess the effect of different control regimes, in terms of population persistence and cost. In particular we consider the effects of each regime on the probability of extinction and the expected time to extinction, and compare the control methods in terms of the expected total cost of each regime over the life of the population. The usefulness of our results will be illustrated with reference to the control of a koala population inhabiting Kangaroo Island, Australia.