Numerical behavior of a zooplankton,phytoplankton and phosphorus system

Several numerical features of a three-dimensional dynamical system which models the three species system made up of phytoplankton, zooplankton and organic phosphorus nutrient in a lake environment are presented. Certain properties of this system, such as the existence of limit cycles, are demonstrated numerically. It is shown that in certain time domains the system may be stiff, thus requiring the use of suitable algorithms.     

富营养化水体中浮游动物对藻类的控制作用

采用室内受控生态系统进行实验.采用于桥水库原水接种后,人工设置营养盐质量浓度梯度,模拟富营养化程度不同的水体,以研究浮游动物的生长情况,及其控制藻类生长的作用.使用SPSS统计软件进行数据分析.结果显示:过高的营养盐浓度,可能会对轮虫的生长产生抑制作用,而甲壳类动物的总数量基本由营养盐质量浓度决定.在四个营养盐质量浓度梯度下,浮游动物与藻类之间均存在一定的显著或极显著相关关系,说明浮游动物均能够发挥一定的控藻作用.当营养盐ρ(TN)=3mg·L-1、ρ(TP)=0.02mg·L-1左右,浮游动物的控藻作用最明显.当营养盐质量浓度过高时,浮游动物的控藻作用受影响.     

Relationship between the zooplankton,phytoplankton, particulate matter and dissolved free amino acids in the Celtic Sea

Estimates of the biomass of zooplankton, phytoplankton and particulate matter collected in the Celtic Sea during mixed-water conditions (on 8 and 9 April 1983) were compared to the concentration and diversity of sixteen dissolved free amino acids (DFAA) measured in seawater and in particles. During a day profile, variations of dissolved amino acids with depth reflected the feeding activity of copepods. The relationship was not apparent in a night profile and other processes, such as heterotrophic utilization of dissolved nitrogen by microorganisms, were thought to be involved. The ratios of total DFAA concentration (nM litre^-1) in the particulate phase over the concentration in seawater ranged from 1 to 200 within the water column. Of the sixteen amino acids measured, ornithine, a decomposition product of arginine, was responsible for more than 70% of the total concentration of DFAA in seawater. In the particles, phenylalanine ranged from 30 to 88% of total DFAA. In seawater this amino acid occurred in the 20 to 40 m depths (1.3 to 9.9% of total DFAA) in the day profile and at 5 m (12.4%) and 80 m (6.4%) in the night profile. Previously it has only been found in very low concentrations (<5%) in seawater, and its presence is considered to be the result of zooplankton feeding.     

Potential effect of phytoplankton colony breakage on the calculation of zooplankton filtration rates

Substantial net production of short chains of the diatom Skeletonema costatum occurred in grazing experiments conducted with Acartia hudsonica (=A. clausi). A general model was then constructed to evaluate the influence of colony breakage during grazing on the calculation of zooplankton filtration and ingestion rates. Breakage can lead to either over-or underestimation of these rates. The magnitude of error is related to breakage probability, particle size distribution, size of the largest colonies, length of the experiment, copepod concentration and actual filtration rates. Size distributions similar to those described in the literature as manifestations of a complex selection for size and particle abundance can be generated by the grazing model under conditions of a uniform filtration rate accompanied by colony breakage. Unless the rate of breakage exceeds the rate of removal and causes net particle production, the occurrence of breakage may go undetected.     

Uptake and release of phosphorus by phytoplankton in the Chesapeake Bay estuary,USA

The phytoplankton uptake and release rates for inorganic phosphate, dissolved organic phosphate and polyphosphate were estimated during 5 cruises on the Chesapeake Bay over a 9-month period. Phosphorus in all pools turned over in several minutes to 100 h, and each soluble pool appeared to contain fractions which were metabolically useful to the phytoplankton. Maximal uptake rates (V _m ) for orthophosphate ranged from 0.02 to 2.95 μg-at P (1.h)^-1 with half saturation constants (K _s ) between 0.09 and 1.72 μg-at P l^-1. At low soluble reactive phosphorus concentrations, the uptake rate of trace ³²P orthophosphate was initially rapid, but declined after 15 to 60 min incubation. The data suggest that the initial uptake phase was dominated by exchange of ³²PO₄ ^≡ for ³¹PO₄ ^≡ in the membrane transport systems whereas the subsequent phase represented the net incorporation of orthophosphate into phytoplankton cells.     

Effects of zooplankton diel vertical migration on a phytoplankton community: A scenario analysis of the underlying mechanisms

A mechanistic model was applied to study the influence of diurnal vertical migration (DVM) of planktonic crustaceans on the succession and composition of the phytoplankton community. While zooplankton was restricted to only one functional group, the phytoplankton community was divided into two functional groups which are distinguished by their maximum growth rates and vulnerability to zooplankton grazing. DVM causes a pulsed grazing regime and may also entail a corresponding reduction of the cumulative daily rates of ingestion and losses of zooplankton. To study the relative importance of these two mechanisms of DVM to phytoplankton we performed a scenario analysis consisting of 5 different scenarios. The results show that DVM has a strong influence on the phytoplankton community. Well edible algae benefit during the first 3–4 weeks of summer stratification by reduced daily grazing. The typical shift from small, well edible algae to larger, poorly or non-edible phytoplankton is distinctly delayed. Under the assumption of unchanged daily grazing, however, a pulsed grazing regime has nearly no influence on the resulting phytoplankton composition. As similar effects are also found for completely non-edible phytoplankton, indirect effects via phosphorus availability must be assumed. Thus, the scenario analysis reveals that the observed effects of DVM on phytoplankton can be explained by a combination of two mechanisms: (1) reduction of the daily zooplankton grazing, and (2) changed assimilation and remineralisation of phosphorus. Surprisingly and in contradiction to earlier reports there is almost no DVM effect on phytoplankton due to the sole action of a pulsed grazing regime.     

Seasonal and areal changes in standing stocks of phytoplankton,zooplankton and micronekton in the eastern tropical pacific

Five standing stocks were measured together at similar latitudes and longitudes on seasonally repetitive cruises in 3 areas — western, eastern, and southern — of the eastern tropical Pacific. The stocks were chlorophyll a at 0 to 150 m depth (mg/m²), night and day zooplankton at 0 to 200 m depth (ml/1000 m³), and night crustacean micronekton and fish-pluscephalopod micronekton at 0 to 200 m depth (ml/1000 m³). The logarithms of the measurements of each stock in each area were subjected to analysis of variance with the following factors: season (2 month period), latitude, and longitude. Seasonal coverage was most comprehensive, with 7 successive periods, in the western area (approximately 16° N to 3° S latitude, 100° to 122° W longitude). Most stocks in most parts of the western area had a simple seasonal cycle of low amplitude, with a single maximum and minimum that usually differed by a factor <2; some stocks in some parts of the area exhibited no seasonal cycle; all statistically significant cycles, except for fish-plus-cephalopod micronekton, were similar in phase. In the other two areas, located broadly to the east and south of the western area, suitable measurements were made at only 2 periods (opposite seasons) of the year. There were indications of phase differences between chlorophyll a and zooplankton in the eastern area, which should be further investigated. Most standing stocks declined gradually from east to west, and were higher in known upwelling areas and areas of shoal thermocline than elsewhere.     

Physioecology of zooplankton. III. Effects of phytoplankton concentration,temperature, and body size on the metabolic rate ofCalanus pacificus

Weight-specific rates of oxygen consumption of actively feeding copepodite stages ofCalanus pacificus Brodsky were measured under various combination of phytoplankton concentration and temperature. The rate decreased logarithmically with a logarithmic increase in dry body weight of copepods, and the relationship between these variables was described using a log-transformed allometric equation. The body-size dependence of the metabolic rate was independent of changes in food concentration and temperature, but the metabolic level increased linearly with a logarithmic increase in temperature and was not significantly affected by changes in food concentration. Respiration rates measured in this study forC. pacificus were about twice as high as rates reported for unfed closely related species of the same genus. An analysis of the metabolic cost of feeding processes suggests that metabolic models derived from feeding models may be of little ecological value at present.Contribution No. 1129 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Description and use of an improved method for determining estuarine zooplankton grazing rates on phytoplankton

A method of rapidly determining zooplankton grazing rates on natural mixed phytoplankton populations using ¹⁴C is described. The method simplifies the design of grazing experiments as the grazing time can be kept short enough to prevent recycling of the isotope, and growth of the phytoplankton substrate. Very high specific activity, ¹⁴C-labelled phytoplankton concentrated either by centrifugation or sieving, may be used either as the sole grazing substrate, or as a tracer in natural mixed phytoplankton. Zooplankton, confined in glass jars at either ambient, or higher than ambient concentrations, are permitted to feed on the phytoplankton for periods of 30 min and 2 h, and are then separated by sieving. The zooplankton community grazing rate, or, if the samples are sorted into species, the individual species grazing rates, can be determined after scintillation counting of the zooplankton. The rate of appearance of ¹⁴C-labelled phytoplankton in the zooplankton is an estimate of the grazing rate, and the slope of the line joining the grazing rates at various phytoplankton concentrations gives an estimate of the grazing rate constant for the zooplankton population. The method provides a quick way of obtaining both zooplankton population, and individual species grazing rates on natural mixed phytoplankton. In two experiments, labelled phytoplankton was used as the sole grazing substrate in concentrations ranging between 0.4 and 5 times ambient levels. Grazing rate constants, for net-caught zooplankton concentrated to 46 times (Experiment 1) and 28 times (Experiment, 2) ambient estuarine levels were-0.14and-0.12 of the phytoplankton standing stock per day, respectively. There was a linear increase in the amount of phytoplankton grazed with an increase in phytoplankton concentration up to four times ambient phytoplankton levels. When tracer amounts of labelled phytoplankton were added to samples containing both phytoplankton and zooplankton at ambient concentrations the grazing rate constants were-0.28 and-0.42 of the phytoplankton standing stock per day. We conclude that zooplankton grazing was the major control factor of phytoplankton population size during October–November 1975 in South West Arm, Port Hacking, near Sydney, Australia.     

Continuous plankton records: relationships between species of phytoplankton and zooplankton in the seasonal cycle

Seasonal fluctuations in abundance and their geographical variations in the north-east Atlantic Ocean and the North Sea for 49 taxa of both phytoplankton and zooplankton have been studied by multivariate techniques. Most of the observed variability in both seasonal and geographical fluctuations in abundance can be attributed to differences between the taxa with respect to population growth rates modulated by temperature coupled with the distribution of overwintering stocks, with the overall pattern also influenced by the timing of the establishment and breakdown of vertical stability in the water column. The observations indicate that for any species, the similarity between its seasonal cycles in the various areas, which represent a wide range of hydrographic regimes, is appreciably greater than any differences induced by interaction between the species in any particular area.     

Physioecology of zooplankton. IV. Effects of phytoplankton concentration,temperature, and body size on the net production efficiency ofCalanus pacificus

Weight-specific rates of individual production, total metabolic expenditure and assimilation, and net production efficiencies were estimated forCalanus pacificus Brodsky of selected body weights cultured at various phytoplankton concentrations and temperatures. The weight-specific rate of individual production increased hyperbolically with food concentration, and the maximum rate of individual production decreased logarithmically with a linear increase in body weight propotionally more at high than at low temperature. The weight-specific rate of total metabolic expenditure decreased logarithmically with increasing body weight and was unaffected by changes in food concentration. The effects of food concentration and temperature on the weight-specific rate of assimilation were similar to those on the rate of individual production, but the effect of body size differed considerably. The diversity in the temperature and body-size dependence of the maximum weight-specific rates of various physiological processes suggest (1) that, except for the metabolic rate, the allometric model (log-log relation) is inadequate for describing relationships between maximum rates of physiological processes and body size within species, and (2) that the common assumption that temperature affects the rates of various physiological processes in similar ways is not justified. Net production efficiency increased hyperbolically with food concentration, and the maximum production efficiency first increased slightly and then decreased gradually with increasing body weight. Small copepods attained higher efficiency at high temperature but larger ones did so at low temperature. The critical food concentrations for production efficiency and for the rate of individual production increased with increasing temperature and body size. Because of the effects of interactions among critical food concentration, temperature, and body size on the rates of growth and individual production and on net production efficiency, early development stages ofC. pacificus optimized growth and food conversion efficiency at high temperature, but late stages, particularly at low food concentrations, grew best and transformed food more efficiently at low temperature.Contribution No. 1130 From the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Optimization of exergy and implications of body sizes of phytoplankton and zooplankton in an aquatic ecosystem model

Size appears to be an important parameter in ecological processes. All physiological processes vary with body size ranging from small microorganisms to higher mammals. In this model, five state variables — phosphorus, detritus, phytoplankton, zooplankton and fish are considered. We study the implications of body sizes of phytoplankton and zooplankton for total system dynamics by optimizing exergy as a goal function for system performance indicator. The rates of different sub-processes of phytoplankton and zooplankton are calculated, by means of allometric relationships of their body sizes. We run the model with different combinations of body sizes of phytoplankton and zooplankton and observe the overall biomass of phytoplankton, zooplankton and fish. The highest exergy values in different combinations of phytoplankton and zooplankton size indicate the maximum biomass of fish with relative proportions of phytoplankton and zooplankton. We also test the effect of phosphorus input conditions corresponding to oligotrophic, mesotrophic, eutrophic system on its dynamics. The average exergy to be maximized over phytoplankton and zooplankton size was computed when the system reached a steady state. Since this state is often a limit cycle, and the exergy copies this behaviour, we averaged the exergy computed for 365 days (duration of 1 year) in the stable period of the run. In mesotrophic condition, maximum fish biomass with relative proportional ratio of phytoplankton, zooplankton is recorded for phytoplankton size class 3.12 (log V μm³ volume) and zooplankton size 4 (log V μm³ volume). In oligotrophic condition the highest average exergy is obtained in between phytoplankton size 1.48 (log V μm³ volume) and zooplankton size 4 (log V μm³ volume), whereas in eutrophic condition the result shows the highest exergy in the combination of phytoplankton size 5.25 (log V μm³ volume) and zooplankton size 4 (log V μm³ volume).     

Evidence for a three-way trade-off between nitrogen and phosphorus competitive abilities and cell size in phytoplankton

Trade-offs among functional traits are essential for explaining community structure and species coexistence. While two-way trade-offs have been investigated in many systems, higher-dimensional trade-offs remain largely hypothetical. Here we demonstrate a three-way trade-off between cell size and competitive abilities for nitrogen and phosphorus in marine and freshwater phytoplankton. At a given cell size, competitive abilities for N and P are negatively correlated, but as cell size increases, competitive ability decreases for both nutrients. The relative importance of the two trade-off axes appears to be environment dependent, suggesting different selective pressures: freshwater phytoplankton separate more along the N vs. P competition axis, and marine phytoplankton separate more along the nutrient competition vs. cell size axis. Our results demonstrate the multidimensional nature of key trade-offs among traits and suggest that such trade-offs may drive species interactions and structure ecological communities.     

富营养化水体降磷对浮游植物群落结构特征的影响

浮游植物是水生态系统中物质循环和能量流动的基础,作为初级生产者,浮游植物的群落结构直接影响着水生态系统的结构和功能。在水产养殖生产中,如何根据养殖生物对生活环境的需求开展精准培水、定向培水,培养养殖生物所需要的浮游植物,在维持养殖水域生态平衡的同时又能为养殖生物提供一定的饵料资源,这一直是摆在水产科技工作者面前的重要难题和研究热点。已有的资料大都是通过添加磷的方式研究磷改变对浮游植物生长的影响,而有关富营养化水体降磷对浮游植物群落结构影响的研究尚未见报道。为此,试验通过向取自富营养化湖泊的水体中加入磷去除剂,采用Pielou均匀度指数、Mcnaughton优势度指数和Shannon多样性指数,研究自然水体中的磷被降低后水体浮游植物群落结构的变化情况。结果表明,所取富营养化水体中共检出绿藻（Chlorophyta）、硅藻（Bacillariophyta）、蓝藻（Cyanophyta）、裸藻（Euglenophyta）、隐藻（Cryptophyta）、甲藻（Pyrrophyta）6门29种（包括变种和变型）;其中绿藻、蓝藻、硅藻、隐藻、裸藻、甲藻分别有7、4、2、1、1种,分别占总种数的24.13%、13.79%、6.90%、3.45%、3.45%。富营养化水体降磷后,虽然试验组和对照组在浮游植物种类组成上没有差异,但浮游植物群落结构特征发生了很大变化,浮游植物数量明显降低,由13 238.8×104cells·L-1降低至3 997.5×104cells·L-1,下降了69.8%;浮游植物优势种从1门（蓝藻（Cyanophyta））6种增加到3门（绿藻（Chlorophyta）、硅藻（Bacillariophyta）、蓝藻（Cyanophyta））12种,优势度指数从97.29%降低至86.30%,优势种门数和优势种种数远远高于对照组,优势度明显低于对照组;同时,浮游植物多样性指数和均匀度分别从1.85和0.38升高至2.60和0.54,显示出试验组浮游植物多样性和均匀度优于对照组。研究表明富营养化水体降磷对浮游植物群落结构产生了明显影响,使群落结构处于更加复杂、完整和稳定的状态。     

Photosynthate partitioning by phytoplankton in a New Zealand coastal upwelling system

A stimulation model of copepod population dynamics (development rate, fecundity, and mortality) was used to compute the predatory consumption necessary to control population growth in three dominant copepod species (Pseudocalanus sp., Paracalanus parvus, and Calanus finmarchicus) on Georges Bank, given observed seasonal cycles of copepod and predator populations. The model also calculated secondary production of each species. Copepod development rate and fecundity were functions of temperature while mortality was a function of predator abundance and consumption rate. Daily inputs of temperature and predator abundance (chaetognaths, ctenophores, and Centropages spp.) were derived from equations fit to field data. Model runs were made with various consumption rates until the model output matched observed copepod seasonal cycles. Computed consumption rates were low compared with published values from field and laboratory studies indicating that, even at conservative estimates of consumption, predators are able to control these copepod populations. Combined annual secondary production by the small copepod species, Pseudocalanus sp. and P. parvus, was nearly twice that of the larger C. finmarchicus with P. parvus having the highest total annual production.     

Physioecology of zooplankton. I. Effects of phytoplankton concentration,temperature, and body size on the growth rate of Calanus pacificus and Pseudocalanus sp.

Changes in dry weight and in weight-specific growth rates were measured for copepodite stages of Calanus pacificus Brodsky and Pseudocalanus sp. cultured under various combinations of phytoplankton concentration and temperature. Mean dry weight of early copepodites was relatively unaffected by either food concentration or temperature, but mean dry weight of late stages increased hyperbolically with food concentration and was inversely related to temperature. The food concentration at which maximum body weight was attained increased with increasing temperature and body size, and it was considerably higher for C. pacificus than for Pseudocalanus sp. This suggests that final body size of small species of copepods may be determined primarily by temperature, whereas final body size of large species may be more dependent on food concentration than on temperature. Individual body weight increased sigmoidally with age. The weight-specific growth rate increased hyperbolically with food concentration. The maximum growth rate decreased logarithmically with a linear increase in body weight, and the slope of the lines was proportional to temperature. The critical food concentration for growth increased with body size proportionally more at high than at low temperature, and it was considerably higher for C. pacificus than for Pseudocalanus sp. Because of these interactions, early copepodites optimized growth at high temperature, even at low food concentrations, but under similar food conditions late stages attained higher growth at low temperature. The same growth patterns were found for both species, but the rates were significantly higher for the larger species, C. pacificus, than for the smaller one, Pseudocalanus sp. On the basis of findings in this study and of analyses of relationships between the maximum growth rate, body size, and temperature from other studies it is postulated (1) that the extrapolation of growth rates from one species to another on the basis of similarity in body size is not justified, even for taxonomically related species; (2) that the allometric model is inadequate for describing the relationship between the maximum weight-specific growth rate and body size at the intraspecific level; (3) that the body-size dependence of this rate is strongly influenced by temperature; and (4) that species of zooplankton seem to be geographically and vertically distributed, in relation to body size and food availability, to optimize growth rates at various stages of their life cycles.Contribution No. 1127 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Nutrient regeneration by zooplankton during a red tide off Peru,with notes on biomass and species composition of zooplankton

During March and April 1976, a red tide, dominated by the dinoflagellate Gymnodinium splendens Lebour, developed in the vicinity of 15°06'S and 75°31'W off Peru. At the height of the bloom, the euphotic zone was 6 m deep and the chlorophyll a at the surface was 48 g l^-1. A daily collection of zooplankton at 09.00 hrs showed large fluctuations of biomass, from 0.2 to 3.84 g dry weight m^-2 in a water column of 120m. Copepodids and nauplii dominated the collections. During a period of reduced wind, the adult copepods were a mixture of the species characteristic of the coastal upwelling system and the neritic species associated with more northerly, tropical waters. Nitrogen regeneration by the zooplankton varied with the development of the bloom, the type of zooplankton dominating the experiment, and biomass fluctuations, but never accounted for more than 25% of the nitrogen uptake by phytoplankton.     

The “planktonkreisel”, a new device for culturing zooplankton

A new device for culturing marine planktonic organisms, the planktonkreisel, is described. It consists of a round glass vessel, a central column facilitating a continuous rotating water current via a jet outlet and an inside sand filter. Its usefulness is documented on the grounds of successful culture and rearing experiments on ctenophores, a chaetognath and meroplanktonic organisms.     

Physioecology of zooplankton. II. Effects of phytoplankton concentration,temperature, and body size on the development and molting rates of Calanus pacificus and Pseudocalanus sp.

Developmental time and stage duration for Calanus pacificus Brodsky and Pseudocalanus sp. and the rate of loss of body carbon by molting for C. pacificus were estimated for copepodite stages cultured under various combinations of phytoplankton concentration and temperature. Mean development time and stage duration for C. pacificus decreased hyperbolically with increasing food concentration, and the minimum time required for reaching a given stage decreased logarithmically with a logarithmic increase in temperature. Low temperature retarded the development of early stages proportionally more than that of late stages, and stage duration increased logarithmically with increasing body weight. Therefore, copepodite development was not isochronal. The rate of loss of body carbon by molting was small, ranging from 0.2 to 2% day^-1. This rate increased hyperbolically with food concentration and was linearly related to the growth rate. The critical food concentration for the rates of development and molting increased with temperature and stage of development, but these rates were less dependent on food concentration than the growth rate. The development rate of Pseudocalanus sp. was higher than that of C. pacificus, and was less influenced by changes in food concentration and temperature. It is postulated that the inverse relationship between temperature and body size results from a differential effect of temperature and body size on the rates of growth and development. That is, with increasing body size the growth rate tends to become temperature-independent, but the development rate remains proportional to temperature. Thus, copepodites growing at low temperature can experience a greater weight increment between molting periods than individuals growing at high temperature, because the growth rate is similar at all temperatures but stage duration is longer at low temperature.Contribution No. 1128 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Small-scale pattern of a California current zooplankton assemblage

The small-scale vertical (10s of meters) and horizontal (100s to 1000s of meters) distribution of zooplankton in the California Current near Guadalupe Island, Baja California, Mexico was studied. Vertical distributions were sampled using a vertically-towed Longhurst-Hardy Plankton Recorder (LHPR) which gave a sequence of samples, each integrated over about 5 m, from 250 m to the surface. Because of the sampling biases of the LHPR, details in vertical structure of less than 15 to 20 m were not considered. The pattern of the vertical tows and the variability of the integrated counts of species were used to infer horizontal distribution. Four series of 8 tows each were taken around noon and midnight over a 2-day period. Four of the 8 tows in each series were randomly positioned within 2000 m of a parachute drogue (first day) or a fixed geographic position (second day); four were replicate tows taken at the drogue or the fixed position. Sixty-seven taxonomic categories were counted. The replicate tows, separated by no more than a few hundred meters, gave more similar vertical profiles for species than did the random tows, with separations of 100s to 1000s of meters. The night replicate tows showed less variability in depth distribution than did either the night random or any of the day tows, leading to the hypothesis that the vertical distributions observed were generated by interactions of the organisms' diel behavior with internal waves. Variability of abundance estimates using the integrated counts was the same for both replicate and random tows, indicating that horizontal patches may be smaller than 100 m. No evidence was found for a day-night change in patch size, or for a consistent overlapping of patches of different species. Replicate tows gave more similar estimates of community structure (relative proportions of species) than did random tows. Overall day community structure was more similar between tows than night structure. Similarities in species' proportions of any random tow to the replicate tows or to other random tows of a series decreased with increasing distance between the tows being compared. This decrease was greater for the night samples, suggesting that community structure is more heterogeneous at night.This study is based in part on a dissertation submitted in partial fulfillment of the requirements for the Ph.D. degree at the University of California, San Diego, and was supported by National Science Foundation Grant GB 12413 and the Marine Life Research Program of the Scripps Institution of Oceanography.