NAVAL POSTGRADUATE SCHOOL Monterey, California THESIS MICROPLANKTONIC ATP-BIOMASS AND GTP-PRODUCTIVITY ASSOCIATED WITH UPWELLING OFF PT. SUR, CALIFORNIA by Sherman Hughes Bronsink September 1980 Thesis Advisor: E. D. Traganza Approved for public release; distribution unlimited T1 97449 UNCLASSIFIED SECUPlTY CLASSIFICATION OF THIS » » r. E fW*on Data Kntarad) fch£13KA»2 REPORT DOCUMENTATION PAGE ' SI^QHT NUWiM READ INSTRUCTIONS BEFORE COMPLETING FORM 2. OOVT ACCESSION MO, 1. RECIPIENT'S CATALOG NUMICH 4 Tl TL £ <*nd isjbtut*) Microplanktonic ATP-Biomass and GTP-Productivity Associated with Upwelling off Pt. Sur, California s. type of report * period covered Master's Thesis; September 1980 «■ PERFORMING ORG. REPORT NUMtIR 7. AUTMORf«J Sherman Hughes Bronsink ». CONTRACT OR GRANT NUMtCRftj t. PERFORMING ORGANIZATION NAME AND AOORESS Naval Postgraduate School Monterey, California 10. PROGRAM ELEMENT. PROJECT, TA&k AREA * WORK UNIT NUMBERS ii CONTROLLING OFFICE NAME ANO ADDRESS Naval Postgraduate School Monterey, California 12. REPORT DATE September 1980 IS. NUMBER OF PAGES 70 TT MONITORING AGENCY NAME A AOOPESSfff dittatant tram Controlling Ottlea) IS. SECURITY CLASS, (of iMt roport) Unclassified 11* DCCLASSIFl CATION/ DOWN GRADING SCHEDULE l«. DISTRIBUTION STATEMENT (at thta Hamart) Approved for public release; distribution unlimited. 17. DISTRIBUTION STATEMENT fat tha aaatraet antatad In BlaaM 30, II dltlarant Xoport) H SUPPLEMENTARY NOTES IS KEY WORDS (Cantlnua on rawaraa alda It naeaaa* Adenosine Triphosphate Guanosine Triphosphate Productivity Biomass Fluxes and tdontlty my aloe* nummar) Phosphate Chlorophyll Temperature Upwelling Satellites Nutrients Nitrate 20 ABSTRACT (Contlnua an tarr an* tdontlty »r alack toot) Microplanktonic ATP-bi in upwelling features off by adenosine triphosphate have a definite preferenti temperature and nutrients Productivity, as sampl triphosphate (GTP) samplin omass and GTP-productivity were studied Pt. Sur, California. Biomass, determined (ATP) and chlorophyll a, was found to al location in the strong gradients of on the equatorward edge of the feature, ed using the new method of guanosine g was found to be high throughout the FORM I JAN 7 J (Page 1) DD 1473 EDITION OF I MOV «S IS OBSOLETE S/N 010J-0 J4- A*01 : UNCLASSIFIED limBftfaiBBiMMItflBf^ tutor ad) UNCLASSIFIED j»cuwT» cl«hihc«tiqw or Tmi >>atfnM n*«« 3-.»~* thermo-nutrient gradient associated with an upwelling feature in an early stage of development. The measurement of productivity and "growth potential" using GTP and GTP/ATP ratios was found to be a valuable tool in examining rapidly changing upwelling features. Preliminary results obtained using the GTP technique suggest a relation- ship between the flux of nutrients across ocean fronts and planktonic productivity associated with frontogenic coastal upwelling systems. DD Form 1473 „, 1 Jan 73 S/N 0102-014-6601 UNCLASSIFIED HtU'lTV CUAHtHCATlQM Qg Tjjjj PiOtfWhmm Dmla t*<»'*4) Approved for public release; distribution unlimited Microplanktonic ATP-Biomass and GTP-Productivity Associated with Upwelling off Pt. Sur, California by Sherman Hughes Bronsink Lieutenant, United States Navy B.S., Western Washington State College, 1973 M.S., University of West Florida, 1975 Submitted in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE IN OCEANOGRAPHY from the NAVAL POSTGRADUATE SCHOOL September 1980 ABSTRACT Microplanktonic ATP-biomass and GTP-productivity were studied in upwelling features off Pt. Sur, California. Bio- mass, determined by adenosine triphosphate (ATP) and chlor- ophyll-a, was found to have a definite preferential location in the strong gradients of temperature and nutrients on the equatorward edge of the feature. Productivity, as sampled using the new method of guanosine triphosphate (GTP) sampling was found to be high throughout the thermo-nutrient gradient associated with an upwelling feature in an early stage of development. The measurement of productivity and "growth potential" using GTP and GTP/ATP ratios was found to be a valuable tool in examining rapidly changing upwelling features. Preliminary results obtained using the GTP technique suggest a relation- ship between the flux of nutrients across ocean fronts and planktonic productivity associated with frontogenic coastal upwelling systems. 4 TABLE OF CONTENTS I. INTRODUCTION 11 A. HORIZONTAL NUTRIENT FLUXES 12 B. VERTICAL NUTRIENT FLUXES 13 C. FRONTAL NUTRIENT FLUXES 14 D. ASSAYING ATP AND GTP 14 E. GUANOSINE TRIPHOSPHATE I7 II. METHODS 18 A. CHLOROPHYLL a 18 B. ATP AND AATP 19 C. AATP VS. GTP CORRELATION EXPERIMENT 21 III. RESULTS 23 A. AATP VS. GTP CORRELATION EXPERIMENT 23 B. CRUISES 24 1. September and November Cruises 24 2. June Cruise 52 IV. DISCUSSION 55 A. AATP VS. GTP CORRELATION EXPERIMENT 55 B. THE JUNE CRUISE 55 V. CONCLUSIONS 62 BIBLIOGRAPHY 6 5 INITIAL DISTRIBUTION LIST 68 LIST OF TABLES I. AATP as a Measure of GTP in a Binucleotide Mixture 22 II. Ranges of ATP, AATP, AATP/ATP, and Chlorophyll a for the June 11-12, 1980 Cruise 53 III. Correlation Coefficients between ATP, AATP, AATP, Chlorophyll a, and Temperature for the June 1980 Cruise 54 LIST OF FIGURES 1. Kinetics of ATP, GTP, UTP , and CTP-dependent light emissions using crude luciferase preparations-- 16 2. Bathymetry of the study area 25 3. Cruise track for the September 1979 cruise 26 4. Nitrate, phosphate, and sea surface temperature versus elapsed distance along the track of the September 27-28, 1980 cruise 27 5. ATP, chlorophyll a, and sea surface temperature versus elapsed distance along the track of the September 27-28, 1980 cruise 29 6. ATP distribution for the September 1979 cruise 31 7. Sea surface temperature for the September 1979 cruise 32 8. Nitrate distribution for the September 1979 cruise 33 9. Phosphate distribution for the September 1979 cruise 34 10. Cruise track for the November 1979 cruise 35 11. Nitrate and sea surface temperature versus elapsed distance along the track of the November 29-30, 1979 cruise 36 12. ATP, chlorophyll a, and sea surface temperature versus elapsed distance along the track of the November 29-30, 1979 cruise 37 13. ATP distribution for the November 1979 cruise 38 14. Sea surface temperature for the November 1979 cruise 39 15. Nitrate distribution for the November 1979 cruise 40 16. Cruise track for the June 1980 cruise 41 17. Nitrate, phosphate, and sea surface temperature versus elapsed distance along the cruise track of the June 10-11, 1980 cruise 42 18. ATP, chlorophyll a, and sea surface temperature versus elapsed distance along the track of the June 1980 cruise 43 19. ATP, AATP/ATP ratio, and sea surface tempera- ture versus elapsed distance along the track of the June 1980 cruise 44 20. ATP distribution for the June 1980 cruise 45 21. AATP/ATP ratio distribution for the June 1980 cruise 46 22. AATP distribution for the June 1980 cruise 47 23. Chlorophyll distribution for the June 1980 cruise 48 24. Sea surface temperature for the June 1980 cruise 4y 25. Nitrate distribution for the June 1980 cruise 50 26. Phosphate distribution for the June 1980 cruise- 51 LIST OF PHOTOGRAPHIC PLATES 1. TIROS-N Satellite IR Image of the California Coast, 9 June 1980 56 ACKNOWLEDGEMENTS This thesis is a result of ongoing research in chemical oceanography at the Naval Postgraduate School which is sup- ported by the Office of Naval Research, Code 482, NORDA, NSTL Station, Bay St. Louis, Mississippi. I thank the sponsors and key individuals whose assistance made this task possible, including: Dr. Eugene D. Traganza, my thesis advisor and principal investigator; Captain W. W. Reynolds and the crew of the R/V ACANIA; Dana Austin and Bonita Hunter of the Naval Postgraduate School. Lastly, I would like to give special thanks to Walter E. Hanson, without whose assistance I would have been unable to complete this thesis . 10 I. INTRODUCTION This thesis examines the possibility that optimal growth conditions for microplankton occur near oceanic fronts as a result of horizontal nutrient fluxes. These fluxes may result from the sharp nutrient and thermal gradients that define such fronts. The research conducted here is part of a continuing study of the oceanic fronts and eddies formed by upwelling off Pt . Sur, California. During previous re- search [Traganza, Nestor, and McDonald, 1980a; Traganza, Conrad, and Breaker, 1980b], comparison of satellite thermal imagery to in situ biomass, nutrient, and thermal measurements showed large spatial variations in biomass concentrations. Microplanktonic blooms appeared greatest on the equatorward edge of the curl of "cyclonic upwelling systems" and adjacent to the sharp gradients in nutrients and temperature. There are several possible reasons for this distribution. The biomass may be advected into the area where it is found. It may be there initially, or it may appear as a preferential growth response to the newly formed front. Preferential growth should be the result of optimal environmental conditions. Since large microplanktonic blooms Gas measured by chlorophyll a and adenosine triphosphate CATP) ) appeared near sharp gradients of nutrients and temper- ature between upwelled, nutrient-rich, biochemically new 11 water and nutrient depleted, biochemically old, oceanic water, a nutrient and thermal flux may provide the neces- sary growth conditions. A horizontal, outward flow from the upwelling system may maintain the gradients ("a natural chemostat") and simultaneously provide a mechanism for biological seeding of fronts with the newly upwelled organ- isms [Traganza, et al . , 1980b]. A. HORIZONTAL NUTRIENT FLUXES Other studies have related phytoplankton production to horizontal and vertical nutrient gradients in Monterey Bay [Lasley, 1977] and to vertical fluxes of nutrients off Southern California [Eppley, Renger, Harrison, 1979]. Lasley [1977] computed the variations of phosphate, nitrate, and chlorophyll a using the distribution of vari- ables equation [Sverdrup, e_t al. , 1942] : 9C „ ac Ar 3C 3C t v 82C D ,, at - "ua7 "VT7 ' wa7 + Ka5^ + R ^ where C represents the concentration of the variable (at 5m in Lasley' s case), t is time, u, v, and w are water velocity in the x, y, and z directions (x defined as parallel to the coast, y perpendicular to the coast, and z as depth), K is the lateral eddy diffusion coefficient. R is a nonconserv- ative term which represents uptake or assimilation of the given nutrient by phytoplankton. 12 From a knowledge of the lateral eddy dif fusitivity coefficient for Monterey Bay, the surface circulation and the nutrient distributions within the bay, it was possible to compute the nutrient fluxes and compare them to the biomass. Measurements of the movement of isopleths of nutrients provided actual flux measurements for comparison. A study of a bay system has several advantages over the study of an exposed coastal upwelling feature interacting with a Western boundary current. The diffusivity coefficient (K) was already known for this area. Surface circulation and nutrient fluxes could be measured with relative ease, since the "feature" (the bay) was a fixed one. The peak in phytoplanktonic biomass (as measured by chlorophyll a) was found near Pt. Pinos where the gradients of nutrients (and presumably the fluxes) were greatest. B. VERTICAL NUTRIENT FLUXES Eppley [1979] postulated that carbon assimilation and therefore productivity could be directly related to vertical nitrate fluxes (dN/dz) following the equation: , K (dN/dz) uP(production • m" ) ^ 2) zo where P is phytoplankton standing stock (mg/m3) , y is its specific growth rate (day" ) at depth z , and depth z (m) n r o v is the depth at which the nitrate concentration has increased 13 to 1 jig - atom • liter" . The depth z is arbitrary and could be chosen for another concentration if the coefficient K were also changed. C, FRONTAL NUTRIENT FLUXES In this study, biomass (ATP and chlorophyll a), nutrients (nitrates and phosphates) , and sea surface temperature were collected on a developing upwelling feature detected by satellite (IR) imagery of the Central California coast on June 10-11, 1980. Productivity and growth potential (in- stantaneous productivity per unit biomass) were examined using the new method of guanosine triphosphate sampling [Karl, 1978]. Linear plots following the ship's track and surface maps of these parameters were constructed to show frontal loca- tions (temperature and nutrients) , concentrations of biomass (based on ATP and chlorophyll a) , areas of high productivity (based on high AATP; see below), and areas of high growth potential (based on high AATP/ATP; see below). D. ASSAYING ATP AND GTP The luciferin- luciferase assay principle for ATP [Holm- Hansen and Booth, 1966; Holm-Hansen and Karl, 1976] has been widely used in recent years. It involves measuring the light emission from the reaction FL ATP + LH2 + 02 > AMP + PPX - C02 + P + hv 3) 14 [Holm-Hansen and Karl, 1978] where LH9 is reduced luciferin, FL is firefly luciferase, PP, is inorganic pyrophosphate, P is product, and hv is light energy. It is relatively easy to extract ATP from "particulates" in sea water with the appropriate solvent. Once extracted, ATP can be measured and used to estimate living carbon biomass by application of a conversion factor representing the mean C/ATP ratio of the microbial (bacteria, algae, and microzooplankton) biomass [Holm-Hansen, 1969]. Even with the right solvent and appropriate analytical procedure there are problems associated with this assay method. The conver- sion factor is somewhat species dependant, and it is depend- ent on the energy charge (physiological state) , and variation of the total adenylate pool, AMP + ADP + ATP [Traganza, 1979]. Diel and other effects have also been noted [Hunter, 1979] . Another problem area concerns the analytical interference of non-adenine nucleotide triphosphates (NTPs) in the assay [Rasmussen and Nielsen, 1968]. When the assay is conducted using crude firefly luciferase and integrated light readings (60 seconds or more of light emissions) , Guanosine tri- phosphate (GTP) , uridine triphosphate (UTP) , and cystosine triphosphate (CTP) all alter the light emission character- istics. The effect is most pronounced when these NTPs are at or near the concentration of ATP. Crystaline luciferase enzyme is ATP specific [McElroy and Green, 1956]. Peak readings (reading the light emission over the first six 15 seconds of the reaction instead of over 60 seconds) mini- mizes the interference, because ATP reacts much more rapidly with luciferin than other NTPs [Figure 1]. iO0 TRIS BUFFER 40 60 80 TIME AFTER INJECTION (sec) Figure 1. Kinetics of ATP, GTP, UTP, and CTP- dependent light emission using crude luciferase preparations. The concentration of each nucleotide was 4 x 10'3 M. [From Karl, 1978]. Measurements of ATP using crude extract and integrated readings result in an overestimat ion of the amount of ATP present because of the presence of other NTPs. This effect can be used to determine GTP levels which, in turn, may prove to be a measurement of productivity. Karl [1979] measured ATP in intertidal sediment extracts using both peak and integrated methods. The difference between these two values (AATP = ATP (integrated) - ATP (peak)) showed a high correlation (r = .96) to the measured amount of GTP present. The AATP values were poorly correlated to any 16 other parameter in the experiment CATP, location the sample was from, etc,)- From this Karl concluded that AATP is useful as a relative measure of GTP, especially in inter- tidal extracts. E. GUANOSINE TRIPHOSPHATE The ability to easily measure GTP is important to the study of mesocale ocean fronts because of its relation to productivity. GTP is known to be essential in the biosyn- thesis of protein. Specifically, it is necessary in the initiation of synthesis, the aminoacyl tRNA binding, and the translocation processes [Watson, 1977] . Since protein synthesis is a major factor in primary productivity, a cor- relation between GTP and primary productivity is expected. Studies have shown a high linear correlation between cellular growth rate and GTP concentration (when normalized to the amount of ATP present, i.e., GTP/ATP) for bacteria [Karl, 1978], algae [Iwamura, et al. , 1963], protozoa [Echetebu and Plesner, 1977], and even fungi [Constantini et al . , 1977]. The ratio of GTP to ATP may represent a "potential growth index" which would be invaluable because it would allow rapid assessment of productivity over large areas. This capability, combined with the rapid measurements of sea surface tempera- ture (SST) and nutrients (nitrate and phosphate) may allow determination of the relation of productivity to fluxes in our rapidly developing systems. 17 II. METHODS Surface values of fluorescence (principally a measure of chlorophyll a) , ATP, and AATP (as an index of relative GTP levels) were obtained off Pt. Sur, California during the development of a "cyclonic upwelling system" [Traganza e_t al ., 1980b] . Temperature and nutrients (nitrate and phosphate) were obtained by Hanson [1980] and used here for interpre- tation of these results. Nutrients (nitrate and phosphate) were sampled at two minute intervals using a Technicon Autoanalyzer . Sea surface temperature (SST) was recorded continuously from an intake located at approximately 2.5 meters depth. Enhanced TIROS-N satellite images were obtained from the National Environmental Satellite Service (NESS) at Redwood City, California. A. CHLOROPHYLL a Fluorescence was measured continuously during the cruises using a Turner Model III fluorometer and converted to pigment concentration (principally chlorophyll a) as described by Lorenzen [1966] . To calibrate the system, discrete 275 ml seawater samples were taken in triplicate every hour. These were filtered through Whatman 4.5 cm diameter G/FC (pore size ca 0.45 pi) glass fiber filters. The filters were folded, placed in polyethelene bags, frozen and analyzed within two 18 weeks, using the methods of Strickland and Parsons [1968]. Chlorophyll a concentrations were determined from the flu- orescence record at two minute intervals to correspond to the nutrient sampling rate. To generate an organic carbon equivalent for comparison with organic carbon derived from ATP, "chlorophyll a" concentrations were converted to carbon units (mg/1) by using the average C/chlorophyll conversion factor of 100 as proposed by Holm-Hansen [1969]. Eppley [1977] has recommended a conversion factor of 54 ±17. The two will be compared in the discussion section. B. ATP AND AATP ATP and AATP samples were taken at approximately 10 min- ute intervals. Fifty ml samples of sea water were filtered through a 200 urn nylon screen and then through a 2.5 cm (diameter) Reeve Angel 984-H (ca. 0.45 ym pore size) glass filter. The filter was then placed in 100°C Trizma (Tris (Hydroxymethyl) aminomethane and hydrochloride) buffer (pH 7.7) to extract the ATP and other nucleotides. The buffer was then frozen for later analysis. The modified ATP method of Holm-Hansen and Karl [1976] was used in the ATP analysis . SAI photometers, Models 1000 and 3000 (SAI, 4060 Sorrento Valley Boulevard, San Diego, California, 92121) were used to determine ATP concentrations from peak height measurements of the light emitted from the samples with an extract of 19 firefly luciferin and luciferase. Comparisons to standards (30, 15, 9, 6, and 3 ng/1) yielded ATP values in ng/1. These were converted to organic carbon biomass (mg/1) by using the average C/ATP conversion factor of 250 proposed by Holm- Hansen [1969] for the "microbial biomass" (bacteria, algae, and microzooplankton) . A relative measure of GTP present was computed using the "AATP" method of Karl [1978]. ATP values were determined using both peak (6 second) and integrated (15 second delay, 60 second assay) methods on an SAI ATP photometer. A strip chart was fitted to the photometer to record peak readings which were converted to ATP concentrations by comparison of peak heights to the peak height of the standards. Integrated readings were given by the photometer's digital readout and converted to ATP concentrations by comparison to the digital readouts of the known standards. The difference between these estimates of the ATP concentration in a given sample (AATP) was used as a relative measure of GTP in the sample. According to Karl, 40 to 50% of the AATP value is GTP (per- sonal communication) . The AATP values were normalized to the ATP by dividing by the amount of ATP present (AATP/ATP) . This ratio provides an index of "potential growth" or an estimate of the instantaneous productivity per unit biomass. 20 C. AATP GTP CORRELATION EXPERIMENT As a test of the validity of using AATP as a measure of GTP on our instruments, a AATP vs. GTP correlation experiment was run using the SAI model 3000 photometer. Known concen- trations of GTP and ATP were mixed in Trizma to represent varying percentages and absolute concentrations which may be found in sea water samples (Table I) . "Peak" and "integrated" values were recorded on a strip chart interfaced to the photometer. 21 TABLE I AATP AS A MEASURE OF GTP IN A BINUCLEOTIDE MIXTURE GTP (ng/1) ATP (ng/1) AATP (CPM*) % GTP (by wt) .3 5.7 7 5 .6 5.4 8 10 .9 8.1 11 10 1.5 4.5 6 25 1.5 13.5 30 10 1.5 28.5 49 5 2.25 6.75 21 25 3.0 3.0 2 50 3.0 27.0 76 10 3.75 11.25 66 25 4. 5 4.5 31 50 7.5 22.5 182 25 7.5 7.5 123 50 9.0 0 0 100 11.25 4.75 137 75 15.0 15.0 269 50 15.0 0.0 79 100 22.5 7.5 329 75 30. 0 286 100 *CPM = Counts Per Minute, proportional to the amount of light emitted in the reaction. Values shown are averages of up to four readings taken at each concentration. 22 III. RESULTS A. AATP VS. GTP CORRELATION EXPERIMENT A linear regression analysis was performed on values of AATP vs. GTP using a WANG Series 700 Advanced Programming Calculator . When all concentrations of ATP and GTP were considered (66 data points), a correlation coefficient or r of .87 was obtained. Adenosine diphosphate (ADP), derived from ATP, is a necessary catalyst for GTP reaction, viz; ADP + GTP -> ATP + GDP 4) ATP ■> ADP + PP + hv 5) with no ATP present initially, the only source of ADP is the crude extract, which limits the reaction. If the values with no ATP present are excluded (leaving 57 data points) , an r value of .92 was obtained. If mixtures with a very low ATP content yielding a AATP of less than 10 counts are also excluded (leaving 36 data points), an r value of .94 was obtained. (In the analysis of the June 10-11, 1980 cruise, only three data points of 189 had a AATP of less than 10 counts. ) 23 B. CRUISES Linear plots of nutrients, temperature and biomass fol- lowing the cruise tracks were computer constructed by an IBM 360. A VERSATEC plotter was used to construct surface contour maps of the parameters in the upwelling features studied. The contour maps were hand smoothed to minimize the effects of advection (the feature moved during the 40 hours that samples were taken which introduced structure into contour maps which did not exist in reality) and computer generated anomalies which occurred at the margins of the data field (the outside edges of the feature were distorted by the boundary conditions set into the computer) . These tract plots and surface maps are presented in Figures 2 to 26. 1 . September and November Cruises Surface maps and linear track maps from the September 27-28, 1979 and November 29-50, 1979 cruises are presented for background (Figures 3 to 15). The November 29-30, 1979 cruise best illustrates the biomass patchiness that was en- countered at the equatorward edge of a feature. The ATP measured biomass shows several "cells" (Figure 6) . Biomass is highest along the equatorward front of the feature where it reaches over 400 ng ATP/1 (100 g C/l) . Other, earlier cruises [Traganza, Conrad, and Greaker, 1980], also showed the highest concentrations of biomass along the equatorward edge of the upwelled feature. Without information on pro- ductivity (or "growth potential" analogous to assimilation 24 BATHYMETRY SCALE 1 750.000 OISTANCES IN KILOMETERS DEPTHS IN METERS 0 121' 30 W CAPE SAN MARTIN Figure 2. Bathymetry of the study area. 25 0053 1620 SHIP'S TRACK 27-28 SEPTEMBER 1979 SCALE 1 : 750.000 DISTANCES IN KILOMETERS TIME : GMT -3d. 10 31. ■ 12 70 -6.35 U.Od 6. i'j 12.70 H.U5 25.40 31. 75 30.10 44 44 50.79 57.14 6 i 49 Figure 3. Cruise track for the September 1979 cruise 26 1 «■ eupseo oiSTiwce L6G 2 LEG 3 TEUPE RATuRE NITRATE PHOSPHATE ELAPSED OiSTANCE km TEMPERATuflE NtTRATE PHOSPHAT6 ELAPSED DISTANCE Figure 4. Nitrate, phosphate, and sea surface temperature versus elapsed distance along the track of the September 27-28, 1980 cruise. 27 TEMPERATURE NITRATE PHOSPHATE N ELAPSED OISTANCE km TEMPERATURE | PHOSPHATE ELAPSED DISTANCE »m Figure 4. Continued 28 ELAPSED DISTANCE 16. LEG 2 LEG 3 •6 *" 14 — TEMPERATURE CHLOROPHYLL 12 10 . -— ELAPSED OISTANCE A TEMPERATURE ELAPSED DISTANCE Figure 5. ATP, chlorophyll a, and sea surface temperature versus elapsed distance along the track of the September 27-28, 1980 cruise. 29 IS LEG 4 LEG 5 L EG 5 LEG a '* /' \ I " —-r- TEMPERATURE CHLOROPHYLL 10 A , -■ - ' ■— * ; V---~- -■ "^~z^- .^ "- ELAPSED DISTANCE r __ "-—-_■ LEG 4 LEG 7 -_— -^ * ' — "^^" — — TEMPERATURE -■■•^ ATP ■ CMLOROPHYIL A /\a A V _^ — - _-^-~^_ -"~~^_ ^^. "^^ -W— — — *T" ELAPSEO DISTANCE Figure 5. Continued. 30 ATP 28 SEPTEMBER 1979 LE 1 750. QOO NCES IN KILOMETERS OURS IN ng/|iter -41.14 -38.10 -31.75 -25.10 -I9.05 -12. 70 -6.35 0.00 6.35 12.70 13.05 23.40 31.75 38.10 11.44 50.79 37.11 Figure 6. ATP distribution for the September 1979 cruise 31 SEA SURFACE TEMPERATURE 27- 28 SEPTEMBER 1979 SCALE 1 : 750.000 OISTANCES IN KILOMETERS CONTOURS IN *C HY.HH -38.10 -31. 75 -2S.H0 -19.05 -12.70 -6.3S 0.00 6.35 12 '0 IS. OS 25. HO 31.75 38.10 itH.UH 50.79 57. 1H 63. H9 Figure 7. Sea surface temperature for the September 1979 cruise. 32 NITRATE 27 • 28 SEPTEMBER 1979 SCALE 1 750,000 DISTANCES IN KILOMETERS CONTOURS IN yuM m.l'l III. MJ II, ,'b t-\ 'in H.nl W.W SO. 79 SMI 63,1'J Figure 8. Nitrate distribution for the September 1979 cruise. .53 PHOSPHATE 27-28 SEPTEMBER 1979 SCALE I ; 7S0.000 DISTANCES IN KILOMETERS CONTOURS IN ^M 121' 30' W \ CAPE SAN MARTIN \1. IJ I'i.OS SS.«ll ll . li I 10 SO. '9 S7. I'l G3.i4d Figure 9. Phosphate distribution for the September 1979 cruise. 34 MONTEREY SHIP'S TRACK 29 - 30 NOVEMBER 1979 SCALE 1 750,000 DISTANCES IN KILOMETERS TIME GMT '21* 30' W CAPE SAN MARTIN Figure 10. Cruise track for the November 1979 cruise S represents a hydro station number. 35 T -rt ■^-V ~'\-'" ' TEMPERATURE NITRATE 1 1 ELAPSED DISTANCE *£MPC BA'Ufic ELAPSED Ot$T*NCE TEMPefl*TUR6 TITRATE .0. ,Lfl 6LAPSEO CSMni F km LEG 10 LEG ii LEG 12 "1 LEG B LEO 9 r~ ■ ■ TEMPERATURE 1 s . ^ V .*-_' "~* N 1 T S A T E — w-y i ■ i i rs """""N ^ i CLiBSEO DISTANCE Figure 11. Nitrate and sea surface temperature versus elapsed distance along the track of the November 29-30, 1979 cruise. 36 II LEG 1 te * — V , . .„— , ^ s " V/vA -wv^^^ v/~" — i — — — — temperature i ■' CHLOBOPHni .0 • 8 •~~*- ^~^ "■ ELAPSED O'STANCE / TC"4PeB4TURE ELAPSED DISTANCE Figure 12. ATP, chlorophyll a, and sea surface temperature versus elapsed distance along the track of the November 29-50, 1979 cruise 37 -44. <44 -38.10 -31.75 -25.110 -J9.05 -12.70 -6.35 0.00 6.35 0 19.05 25.40 31.75 33.10 mj.ti<4 50.79 57. W 63. Figure 15. ATP distribution for the November 1979 cruise. 38 SEA SURFACE TEMPERATURE 29-30 NOVEMBER 1979 SCALE 1 : 750,000 DISTANCES IN KILOMETERS CONTOURS IN "C 121" 30' W I \ CAPE ISAN MARTIN > '5 -25.4Q -:3.:S Figure 14. Sea surface temperature for the November 1979 cruise. 39 I I I NITRATE 29-30 NOVEMBER 1979 SCALE I 750.000 DISTANCES IN KILOMETERS CONTOURS IN «M 121" 30 W \ CAPE -SAN M A R T I •'' ■"'' "•10 il. 's .■>', . w" "TVTos Figure 15. Nitrate distribution for the November 1979 cruise 40 SHIP'S TRACK 10 -11 JUNE 1980 SCALE 1 750,000 DISTANCES IN KILOMETERS TIME : GMT It. vi M.IO VO I 9. (IS 31 . 75 38 10 SO. 79 57. W t3.«9 Figure 16. Cruise track for the June 1980 cruise. 41 ELA»«CO DISTANCE -€M9€R4TURe XTv S. '•; i - .\ .+., .x v .' v. ^y / ^ \ ■. r~i EL4PS60 QlSTAWCt ~N*' ELAPSED DtSTA* :i 1 — TEMPtaiTuftE NITRATE 1 ~~ PHOSPHATE Figure 17. Nitrate, phosphate, and sea surface temperature versus elapsed distance along the track of the June 10-11, 1980 cruise. 42 IB LEG 1 LEG 2 18 - -■— — TEMPERATURE ■*-**»* ATP CHLOROPHYLL m M- 3 r*s < 2 '2 S ^■~~^~ S— -— -— ,. \y . — . r 'v 10 ^v *^ ^^ — --T.. . " :^-— =. — — =1 '..-,--.--._ __-^-^— =— cc£ ^^~ ELAPSED OISIANC TEMPERATURE --'-V ELAPSED DISTANCE Figure 18. ATP, chlorophyll a, and sea surface temperature versus elapsed distance along the track of the June 10-11, 1980 cruise. 43 Carbon &A C«fBOf» flATP ELAPSED DISTANCE LEG t LEG 9 LEL 10 LEG 11 --. 1 • tEMPERftTUflE L . i*fP/ATP 8. "~ : • - - V. - " *s r ;; • -. / .' . "*, ' * - , ' .. EtiPStO GlSTANCL :E mperatube :: APSED DISTANCE Figure 19. ATP, AATP/ATP ratio, and sea surface temperature versus elapsed distance along the track of the June 1980 cruise. 44 ATP 10 -11 JUNE 1980 SCALE 1 : 750,000 DISTANCES IN KILOMETERS CONTOURS IN ng/llter -44.44 -38.10 -31. 7S -25. 40 -19.05 -12.70 -6.35 0.00 6.35 12.70 19.05 25.10 31.75 38.10 44. 44 50.79 57.14 63.49 Figure 20, ATP distribution for the June 1980 cruise. 45 AATP/ATP 10 -11 JUNE 1980 SCALE 1 : 750,000 DISTANCES IN KILOMETERS -HU.1U -38.10 -31.75 -C5H0 -19.05 -15.70 -6.35 0.00 6.35 12.70 19.05 25.10 31.75 38.10 Hl.m 50.79 57. HI Figure 21. AATP/ATP ratio distribution for the June 1980 cruise. 46 -38.10 -31.75 -25.10 -19. OS -12.70 -6.35 0.00 6.35 13. 70 19.05 25.140 31.75 36.10 11. 11 50.79 57. Figure 22. AATP distribution for the June 1980 cruise. 47 -38.10 -31.75 -25.40 -19.05 -15.70 -6.35 0.00 6.35 12. 70 19. OS 25. .10 31.75 50.79 57. in Figure 23. Chlorophyll distribution for the June 1980 cruise. 48 "I / SEA SURFACE TEMPERATURE 10 -11 JUNE 1980 SCALE I 750.000 DISTANCES IN KILOMETERS CONTOURS iN 'C 121' 30 W |X\ CAPE '"\ \ \ \JSAN MARTIN V- \ Figure 24. Sea surface temperature for the June 1980 cruise. 49 ■Eh NITRATE 10 -11 JUNE 1980 SCALE 1 : 750.000 DISTANCES IN KILOMETERS CONTOURS IN yuM \1 21° 30' W \\1 \ A^M CAPE \\\SAN MARTIN Figure 25. Nitrate distribution for the June 1980 cruise. 50 I ' / / ! ' - MONTEREY Ml \ \ 1.6 .' a ., ' \ 1 I J_ V V N •« .8 *? v s S > \ PHOSPHATE 10 -11 JUNE 1980 SCALE 1 : 750.000 DISTANCES IN KILOMETERS CONTOURS IN ^M ».ug -38. ;0 -3!.^S -??.U0 -19.CS -12. 73 -6.33 0.0-: 1?. 70 '.9. OS Jv.YO Figure 26. Phosphate distribution for the June 1980 cruise. 51 number - see comments below)) and surface circulation, the reason for this patchiness could not be considered. 2 . The June Cruise On June 10-11, 1980, in addition to temperature, nutrient, and ATP data, "chlorophyll a" data was collected and, for the first time, AATP and AATP/ATP values were com- puted. The range of these values, which were computed at approximately 10 minute intervals, are given in Table II along with chlorophyll a and ATP. There were 189 ATP, AATP, and AATP/ATP values on the June cruise. 52 TABLE II RANGES OF ATP, AATP, AATP/ATP AND CHLOROPHYLL a FOR THE JUNE 11-12, 1980 CRUISE high low mean standard deviation ATP (ng/1) 1549 48 278 ± 195 AATP (ng/1) 358 .6 83 ± 30 AATP/ATP .57 -.01 .30 ± .11 Chlorophyll a (mg/m ) 5.59 .17 1.11 ± .83 Correlation coefficients for various parameters were computed and are summarized in Table III. 53 TABLE III CORRELATION COEFFICIENTS BETWEEN ATP, AATP, AATP/ATP, CHLOROPHYLL^, AND TEMPERATURE FOR THE JUNE 198 0 CRUISE Correlation of Coefficient ATP to CHL .536 ATP to AATP/ATP .249 ATP to AATP .851 AATP/ATP to CHL .35 AATP/ATP to TEMP - .24 9 AATP/ATP to ATP .613 AATP to TEMP - .218 AATP to CHL .559 54 IV. DISCUSSION A. AATPVS. GTP CORRELATION EXPERIMENT The high correlation of AATP to GTP shows that in a binucleotid solution AATP may be taken as a measure of the relative amount of GTP present. In field samples there are other NTPs present (most importantly UTP) which enter into the reaction. Fortunately, for a given "system" (one with a relatively constant mix of microbial species) the percent of the AATP value which is GTP may be assumed to be constant [Karl, personal communication]. This constant, as mentioned previously, is on the order of 40 to 50% of the signal. B. THE JUNE CRUISE The June cruise shows a feature in what appears to be the "initiation phase" [Conrad, 1980] . The thermal and nutrient gradients are strong and well defined, and there are strong correlations between nitrate and phosphate (r=.96), nitrate and temperature (r=-.96), and phosphate to temperature (r=-.92). These all point to "biochemically new" water [Traganza, et al. , 1980] which has not had its initial conditions (temperature and nutrient concentrations) significantly altered by dynamical and biological processes. The satellite imagery (Plate 1) also shows a thermal pattern which has the appearance of an upwelling feature which is just forming and beginning to develop a cyclonic curl. 55 &S^-dKCMM«Si->s->>C"-: * " ^>;f-iS*,>M$,,i*V!}r ■''■:■' >-"'■■ i-'V""-'*? ■'•■'■■■■' r- : |W #5 :', .^fV'.4--^? ; ' --: is „ ^ . 1 ^4.^^^i^*l^ 8 1 ~*i& Si lit & ***•* Plate 1. TIROS-N satellite IR image o£ the California coast, 9 June 1980. 56 ATP values are high (above 600 ng/1) in two areas (Fig. 20), the area of Monterey Bay off Pt . Pinos described by Lasley [1977J and discussed in the introduction, and in a narrow band along the seaward edge of the feature off Pt. Sur (stretching from about (5, -3) to (-3, 9) on the contour grid) . This band closely follows the intermediate tempera- ture band paralleling the peripheral margin of the satellite inferred feature and is in an area of water which is from 10.5 to 11.5 °Centigrade. This relation of the biomass to intermediate temperatures can also be seen in the linear plots (Figure 18). The maximum ATP concentrations are on the equatorward edge of the developing cyclonic curl (com- pare the SST figure and the ATP figure) which appears to be developing over the Sur Canyon. The nutrients are of inter- mediate concentration. Nitrates vary from 8 to 15 yM and phosphates from 1.2 to 1.8 uM in this area. The feature appears to be beginning a cyclonic spinup which would further concentrate and move this biomass along the equatorward edge of the feature. Chlorophyll shows essentially the same distribution (Figure 23) with the exception of a large "cell" centered about (-1, 6) on the contour grid. Values here reach 3.5 mg/m . There are several possible explanations for this anomaly. ATP is subject to several variations that chloro- phyll a is not. It is energy charge dependent and shows more diel variation than chlorophyll a does [Hunter, 1979]. 57 In addition, the difference may come from sampling technique. The continuous flow of water through the fluorometer ("chlor- ophyll a" determination) is not prefiltered because this produces a clogging problem. This could result in phyto- plankton (i.e., chain formers) larger than 200 microns being excluded from the ATP samples which are prefiltered, but not from the chlorophyll samples. AATP (or "relative GTP") was highly correlated to ATP (r = .85). Since they are not measures of the same parameter (GTP 'v instantaneous growth rate; ATP *v< living biomass) , it can be concluded that all of the biomass is in a relatively uniform productive state. Cellular alkaline phosphatase may interfere with these field measurements. Alkaline phosphatase acts by selectively consuming ATP and GTP. ATP is consumed at a more rapid rate than GTP, destroying evidence of the initial relationship [Karl, 1980] . The results in our field measurements appeared to be too consistent with temperature and nutrient data to have been effected by this problem. However, a separate test as described by Karl [1980 ] has been conducted on our stored field samples by Dana Austin at the Naval Postgraduate School. There are no indications of alkaline phosphatase interference in six samples selected from representative locations. The AATP/ATP ratio or "growth potential" provides infor- mation on the productivity state of the biomass. The "growth potential" is analogous to assimilation number 58 (mg C uptake/mg Chl/hr) . The standard deviation of the AATP/ATP ratio is only 37% of the mean value. This compares to a standard deviation of 70% of the mean for the ATP and 75% of the mean for chlorophyll. In other words, the "growth potential" varied very little and therefore the biomass adjacent to the fronts is in a relatively uniform productivity state. All of the biomass that is located adjacent to the nutrient fronts is present because of real growth. If the biomass were advected into the area, it should not remain at a high, uniform level. The only log- ical conclusion is that the biomass must be present as the result of real growth, i.e., presumably a preferential growth response to a horizontal nutrient flux across the sharp nutrient gradients. The distribution of AATP/ATP is also of interest. The "growth potential" is high in the same area of Monterey Bay off Pt. Pinos described by Lasley [1977] and in the upwelling feature observed off Pt. Sur . In the feature, the "growth potential" is highest just seaward of the biomass peak. This suggests that the middle of the biomass peak is less active than the edges. This is not unexpected. Areas of high biomass are self - limiting in terms of growth, since they often encounter a growth limit such as light limiting or a nutrient shortage (although our data does not show a nutrient shortage). This also points to real growth in the system 59 Cat least in this early stage) because the distribution is as expected for a growing system. The relationship of the biomass and growth potential to the front suggests that the gradients are an important part of the productivity. As mentioned previously, other authors [Lasley, 1977; Eppley, e_t al . , 1979] have related biomass to gradients by considering the biomass and productivity to flux relationships. Both of these authors were able to infer the circulation to compute the flux from the gradients, something that we have been unable to do at this time. The dynamic features we are studying do not lend themselves to the techniques they employed. Lasley used a fixed feature (Monterey Bay) which had known circulation (from current studies and drifters) . Eppley worked with vertical fluxes and was able to calculate the circulation by applying con- tinuity equations. If the surface circulation were better known in upwelled features as described here, the possibility is strong that a relationship between nutrient fluxes and the productivity would be evident. The results would prob- ably be similar to Lasley' s, if the eddy diffusion term and the advection term parallel to the nutrient contours are negligible. This leaves a distribution of variables equation of: at " "U3x- " W 81 + R V This equation relates the changes in concentration of the 60 parameter (nutrients in this case) directly to the fluxes associated with the vertical and horizontal gradients (with the exception, of course, of the biological term, R) . If this equation is accepted as representing the behavior of the nutrients in the feature, there is a good qualitative agreement between the location of the biomass and produc- tivity vs. the fluxes. 61 V. CONCLUSIONS 1. The study of AATP and AATP/ATP ratios can provide valuable insight into productivity and growth potential. This method is especially valuable in studying dynamic features such as fronts and eddies, where a rapidly changing mesoscale environment and the need for large numbers of samples make more traditional methods unworkable. 2. Several conclusions about the initial or early phase of upwelling features can be drawn from the study of the June cruise. a. Areas of high biomass are areas of high produc- tivity. The biomass present appears to be there as the result of optimal growth conditions, not advection. b. The highest growth is in the area surrounding the highest biomass, i.e., intermediate concentrations of biomass appear to be optimal. Even at this early stage in the feature, biomass is beginning to be limited in its growth by some factor. c. Since both high biomass and high growth potential are found in the area of large nutrient and temperature grad- ients, the gradients must have a favorable impact on growth. A nutrient flux, as postulated earlier, may serve to connect the two. Unfortunately, until the surface circulation is better known, computation of surface fluxes is not possible. 62 If equation 6) is applicable to the June feature, however, the good correlation between productivity and fluxes is suggested qualitatively in the data if the velocities across the gradients are relatively constant. d. With the exception of one chlorophyll a "cell" described earlier, the surface maps of ATP and chlorophyll a are essentially the same. A comparison of the surface maps and mean values suggests that the standard conversion ratios of C:ATP (250:1) and C:Chl (100:1) are not entirely accurate in this feature. If a C:Chl ratio of 60:1 were used instead of 100:1, the values would be in closer agreement, suggest- ing that the populations are largely phytoplankton . Eppley [1977] found the ratio of 54 ±17 for C:Chl in phytoplankton populations off Southern California which supports this suggestion. Without specific knowledge of C:Chl and C:ATP ratios in the populations under investigation, one cannot conclude that microzooplankton or bacterial populations are indicated rather than phytoplankton when ATP is larger than chlorophyll . e. Satellite imagery may be used to infer potential biomass location and growth potential, at least in an initial stage of a feature's development but not all features may support biological activity. Studies of features in later stages of development may provide insight into the biomass and growth potential of older features. Applications of this technology may lead to the ability to remotely predict 63 productive regions for fisheries and other applications. Such predictions would be dependent on the source water depth (a function of the feature's age, the season, and the recent wind history [Hanson, 1980]). 64 BIBLIOGRAPHY Conrad, J.W., Relationships Between the Sea Surface Temper- ature and Nutrients in Satellite Detected Oceanic Fronts, M.S. Thesis, Naval Postgraduate School, 1980. Constantini, M.G., Zippel, R. , and Sturani, E. , Biochimica et Biophysica Acta, V. 476, p. 272-278, 1977. Echetebu, CO. , and Plesner, P. , "The Pool of Ribonucleotide Triphosphates in Synchronized Tetrahymena pyrif ormis , " Journal of General Microbiology, V. 103, p. 389-392, 1977. Eppley, R.W. , Harrison, W.G., Chisholm, S.W., and Stewart, E., "particulate Organic Matter in Surface Waters of Southern California and Its Relationship to Phytoplankton," Journal of Marine Resources, V. 35, p. 671-696, 1977. Eppley, R.W., Renger, E.H., and Harrison, W.G., "Nitrate and Phytoplankton Production in Southern California Coastal Waters," Limnology and Oceanography, V. 24, p. 483-494, 1979. Hanson, W.E., Nutrient Study of Mesoscale Thermal Features off Pt. Sur, California, M.S. Thesis, Naval Postgraduate School, 1980. Holm-Hansen, 0. and Boothe, C.R., "The Measurement of ATP in the Ocean and Its Ecological Significance," Limnology and Oceanography, V. 11, p. 510-519, 1966. Holm-Hansen, 0. and Karl, D.M., "Biomass and Adenylate Energy Charge Determination in Microbial Cell Extracts and Environmental Samples," Methods in Enzymology, V. LVII, p. 73-85, 1978. Holm-Hansen, 0. and Karl, D.M. , "Effects of Luciferin Concentration on Quantitative Assay of ATP Using Crude Luciferase Preparations," Analytical Biochemistry, V. 75, p. 100-112, 1976. Holm-Hansen, 0. , "Determination of Microbial Biomass in Ocean Profiles," Limnology and Oceanography, V. 14, p. 740-747, 1969. 65 Hunter, B.L., ATP/Carbon Ratios as a Measure of Phytoplankton Biomass and Productivity, M.S. Thesis , University of Hawaii, 1979. Iwamura, T. , Kanuzawa, T., and Kunazawa, K. , Studies on Micro- algae and Photosynthet ic Bacteria, Japanese Society of Plant Psysiologists , University of Tokyo Press, p. 587-598, 1965. Karl, D.M. , "Occurrence and Ecological Significance of GTP in the Ocean and in Microbial Cells," Applied and Environmental Microbiology, p. 349-355, 1978. Karl, D.M. , "Adenosine Triphosphate and Guanosine Triphosphate Determinations in Intertidal Sediments," Methodology for Biomass Determinations and Microbial Activities in Sedi- ments, ASTM STP 675, CD. Littlefield and P.L. Seyfried, Eds . , American Society for Testing and Materials, p. 5-20, 1979. Karl, D.M. , and Craven, D.B., "Effects of Alkaline Phosphatase Activity on Nucleotide Measurements in Aquatic Microbial Communities," Applied and Environmental Microbiology, V. 40, No. 3, September 1980. Lasley, S.R., Nearshore Circulation and Aging of Water in Monterey Bay, M.A. Thesis, San Jose State University, 1977. Lorenzen, C.J., "A xMethod for the Continuous Measurement of In. Vivo Chlorophyll Concentration," Deep Sea Research, V. 13, p. 223-227, 1966. McElroy, W.D., and Green, A., Archives for Biochemistry and Biophysics, V. 64, p. 257-271, 1956. Rasmussen, H. and Nielsen, R. , Acta Chemica Scandinavia, V. 22, p. 1745-1762, 1968. Strickland, J.D.H., and Parsons, T.R., A Practical Handbook of Seawater Analysis, J.C. Stevenson (Ed. ) , Queen' s Printer and Comptroller of Stationary, p. 49-62 and 71-76, 1968. Sverdrup, H.U., Johnson, M.W., and Fleming, R.H., The Oceans, Their Physics, Chemistry, and General Biology, Prentice Hall, 1942. Traganza, E.D., Naval Postgraduate School Technical Report No. 68-79-006, "Equilibrium Calculations of ATP as a Theoretical Basis for the 'Normal' Range of C/ATP Ratios in Marine Zooplankton," September, 1979. 66 Traganza, E.D., Nestor, D.A. , and McDonald, A.K., "Satellite Observations of a Nutrient Upwelling off the Coast of California," Journal of Geophysical Research, V. 85 No. C7, p. 4101-4106, July 1980. Traganza, E.D., Conrad, J.W., Breaker, L.C., "Satellite Observations of a 'Cyclonic Upwelling System' and 'Giant Plume' in the California Current," Coastal Upwelling, American Geophysical Union (in press), 1980. Watson, J.D., Molecular Biology of the Gene, W.A. Benjamin, Inc. , p. 331-338, 1977. 67 INITIAL DISTRIBUTION LIST No. Copies 1. Defense Technical Information Center 2 Cameron Station Alexandria, VA 22314 2. Library, Code 0142 2 Naval Postgraduate School Monterey, CA 93940 3. Chairman, Code 68 Mr 1 Department of Oceanography Naval Postgraduate School Monterey, CA 93940 4. Chairman, Code 63 Ha 1 Department of Meteorology Naval Postgraduate School Monterey, CA 93940 5. Dr. E. D. Traganza, Code 68 Tg 5 Department of Oceanography Naval Postgraduate School Monterey, CA 93940 6. LT. S. H. Bronsink 5 Helsuppron Three NAS North Island San Diego, CA 92135 7. Director 1 Naval Oceanography Division Navy Observatory 34th § Massachusetts Avenue NW Washington, D.C. 20390 8. Commander 1 Naval Oceanography Command NSTL Station Bay St. Louis, MS 39529 9. Commanding Officer 1 Naval Oceanographic Office NSTL Station Bay St. Louis, MS 59529 68 10. Commanding Officer Fleet Numerical Oceanography Center Monterey, CA 93940 11. Commanding Officer Naval Ocean Research § Development Activity NSTL Station Bay St. Louis, MS 39529 12. Office of Naval Research (Code 482) Naval Ocean Research § Development Activity NSTL Station Bay St. Louis, MS 39529 13. Scientific Liaison Office Office of Naval Research Scripps Institution of Oceanography La Jolla, CA 92037 14. Library Scripps Institution of Oceanography P.O. Box 2367 La Jolla, CA 92037 15. Library Department of Oceanography University of Washington Seattle, WA 98105 16. Library CICESE P.O. Box 4803 San Ysidro, CA 92073 17. Library School of Oceanography Oregon State University Corvallis, OR 97331 18. Commander Oceanographic Systems Pacific Box 1390 Pearl Harbor, HI 96860 19. Chief, Ocean Services Division National Oceanic and Atmospheric Administration 8060 Thirteenth Street Silver Springs, MD 20910 69 20. Chairman, Oceanography Department U.S. Naval Academy- Annapolis, MD 21402 21. Director Naval Oceanography Division (OP952) Navy Department Washington, D.C., 20350 22. Mr. Ben Cagle Office of Naval Research Branch Office 1030 East Green Street Pasadena, CA 91106 23. Dr. Robert E. Stevenson Scientific Liaison Office, ONR Scripps Institution of Oceanography La Jolla, CA 92037 24. Mr. Jerry Norton, Code 68 Department of Oceanography Naval Postgraduate School Monterey, CA 93940 25. Ms. Bonita Hunter, Code 68 Department of Oceanography Naval Postgraduate School Monterey, CA 93940 26. Mr. Laurence Breaker National Environmental Satellite Service 660 Price Avenue Redwood City, CA 94063 27. LT. Carol Jori, Code 68 Department of Oceanography Naval Postgraduate School Monterey, CA 93940 28. LT. Walter Hanson USCG Oceanographic Unit, Bldg. 159-E Navy Yard Annex Washington, D.C. 20595 70 Thesis 190971 B809239 Bronsink c.l Microplanktonic ATP- biomass and GTP-pro- ductivity associated with upwel 1 ing off Pt. Sur, California. 1 The s i s B8C9239 c.l R ., 190971 Bronsink Microplanktonic ATP- biomass and GTP-pro- ductivity associated with upwel 1 ing off Pt. Sur, California.