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UBC Theses and Dissertations

Biological availability of metals in Juan de Fuca Strait Cave, William Rutherford 1977

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THE BIOLOGICAL AVAILABILITY OF METALS IN JUAN DE FUCA STRAIT by WILLIAM RUTHERFORD CAVE B.S c , Un i v e r s i t y of B r i t i s h Columbia, 1975 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE i n THE FACULTY OF GRADUATE STUDIES Department of Zoology We accept t h i s thesis as conforming to the required standards William Rutherford Cave, 1977 THE UNIVERSITY OF BRITISH COLUMBIA A p r i l , 1977 In presenting th is thes is in p a r t i a l fu l f i lment of the requirements f o r an advanced degree at the Un ivers i ty of B r i t i s h Columbia, I agree that the L ibrary sha l l make it f ree ly ava i l ab le f o r reference and study. I fur ther agree that permission for extensive copying of t h i s t h e s i s for scho la r ly purposes may be granted by the Head of my Department or by h is representat ives . It is understood that copying or p u b l i c a t i o n of th is thes is f o r f i nanc ia l gain sha l l not be allowed without my writ ten permission. Department of The Univers i ty of B r i t i s h Columbia 2075 Wesbrook Place Vancouver, Canada V6T 1W5 Pareuchaeta elongata Esterley i i ABSTRACT A r e l a t i o n s h i p between the b i o l o g i c a l a v a i l a b i l i t y of metal i n Juan de Fuca S t r a i t and the presence of organic complexing agents has been suggested. Research c a r r i e d out i n Juan de Fuca S t r a i t during February - June, 1976 showed a seasonal occurrence of increased DOC was d i r e c t l y r e l a t e d to the appearance of higher s a l i n i t y , lower temperature and oxygen, upwelled water and increased surface p r o d u c t i v i t y . i n the S t r a i t . The suggested mechanism f o r upwelling being the W-NW winds which predominate the l a t e Spring-Summer period annually. The b i o l o g i c a l a v a i l a b i l i t y of metals i s reduced i n oceanic water on i n t r o d u c t i o n of upwelled water containing o r g a n i c a l l y complexed metal. The s u r v i v a l of organisms resident i n that water i s reduced while subse-quent 'ageing' of the water may lead to a gradual increase i n the s u r v i v a l . i i i TABLE OF CONTENTS Page TITLE PAGE i ABSTRACT TABLE OF CONTENTS LIST OF TABLES LIST OF FIGURES ACKNOWLEDGMENTS INTRODUCTION 1 MATERIALS AND METHODS 15 i ) In the F i e l d 15 i i ) Bioassays 15 i i i ) DOC Analysis 20 iv) Metal Analysis 21 v) Wind Data 23 vi ) Fraser River 23 v i i ) S t a t i s t i c a l Analysis 24 RESULTS 25 i ) Bioassays, Hydrography, and DOC and Metal Analysis 25 i i ) Wind 40 i i i ) Fraser River Runoff 46 iv) S t a t i s t i c a l Analysis 46 DISCUSSION 54 BIBLIOGRAPHY . 62 APPENDIX 69 0 i v LIST OF TABLES Page TABLE I. Station l o c a t i o n s , cruise numbers and dates f or the cruise on which hydrographic data, bioassay organisms and water were c o l l e c t e d 16 TABLE I I . TABLE I I I . TABLE IV. Percent.survival of the pre-feeding stages of P. selongata E s t e r l e y i n seawater c o l -l e c t e d from 150 m depth at Juan de Fuca Station 7-3 (48° 17.8'N, 124° 06.4'W) 28 Combinations of the bioassay seri e s used to examine the b i o l o g i c a l a v a i l a b i l i t y of metals i n Juan de Fuca S t r a i t .... 29 Background l e v e l s of DOC (Dissolved Or-ganic Carbon) and Metals (Copper, Zinc, Iron and Manganese) i n Seawater c o l l e c t e d from 150 m depth at Juan de Fuca Station 7-3 (48° 17.8'N, 124° 06.4'W) 32 TABLE V. S a l i n i t y , temperature, and oxygen values of the bioassay control water obtained from 150 m at JF7-3. 33 TABLE VI. Gives the percent of the wind which blew i n each d i r e c t i o n per month for the period June 1975 - June 1976. 41 TABLE VII. TABLE VIII. TABLE IX. TABLE X. Gives the c o r r e l a t i o n c o e f f i c i e n t s (r) obtained by c o r r e l a t i o n analysis of v a r i -ables using the UBC B i o l o g i c a l Computing Center Programme CCOEF. 49 Gives the p r o b a b i l i t y of r e j e c t i o n of the n u l l hypothesis (H Q)> with respect to the c o r r e l a t i o n of the v a r i a b l e s , as derived from a table of the d i s t r i b u t i o n of ' t ' (Snedecor ^ 1956) 50 Provides a key to the va r i a b l e name abbre-v i a t i o n s used i n the s t a t i s t i c a l analysis of the r e s u l t s .... 52 A summary of the s i g n i f i c a n t c o r r e l a t i o n s . .. 53 V LIST OF FIGURES Page FIGURE 1. Juan de Fuca S t r a i t and surrounding water showing the positions of JF7-3, GEO 1748 and Cape Beale .... 4 FIGURE 2. Average speed and d i r e c t i o n of monthly winds f o r the period 1961-1963 (Barnes e_t a l . , 1972). 7 FIGURE 3. Shows the major ocean currents of the Northeast P a c i f i c Ocean (Gardner, 1976). .... 9 FIGURE 4. Location of releases, hypothetical paths, and approximate recovery pos i t i o n s of sea-bed d r i f t e r s found between the Columbia River and Vancouver Island, more than 100 km from release point (Barnes e_t a l . , 1972) 11 FIGURE 5. Naupliar stages of Pareuchaeta elongata Este r l e y . .... 18 FIGURE 6. Graphs showing s u r v i v a l s of EDTA addition se r i e s i n con t r o l and UV photo-oxidized sea-water for Feb.-June, 1976. .... 30 FIGURE 7. Graphs showing s u r v i v a l s of metal addition s e r i e s i n control water f or Feb.-June, 1976 34 FIGURE 8. S a l i n i t y , temperature and oxygen f or Feb.-June, 1976 at JF7-3 (48° 17.8'N, 124° 06.3%). 36 FIGURE 9. Gives the wind d i r e c t i o n s (percent of month-l y t o t a l ) experienced at Cape Beale f or the period June 1975 - June 1976. 42 FIGURE 10. Gives the calculated Ekman Transport values for the 5° grids about 47°N, 126 W and 49°N 128°W f or June 1975 - June 1976 44 FIGURE 11. Shows the monthly Fraser River discharge at Hope, B.C. i n cubic meters per second (X 10^)... 47 FIGURE 12a. Mean monthly values of the "upwelling index" for the 20 year period, 1948-1967. 56 FIGURE 12b. Shows a graphic demonstration of the "upwell-ing index" (above, F i g . 12a). Mean monthly values of the computed upwelling indices f o r the 20 year period 1948-1967 (Bakun, 1973) 56 v i ACKNOWLEDGEMENTS I wish to thank Dr. A.G. Lewis for the encouragement, guidance and assistance he has given me during these past two years. Drs. M. Barnes, K.J.F. H a l l , and P.B. Crean also gave valuable assistance i n both my research and i n the preparation of t h i s t h e s i s . I extend my thanks to those who as s i s t e d me with my f i e l d work, Mr. A. Ramnarine, Mr. A. F u l l e r , Mr. S. L i p i n , and Mr. B. Smith, and to the o f f i c e r s and crew of the Canadian Hydrographic Service Ship "Vector" who were outstanding i n t h e i r cooperation and helpfulness. My thanks also to Mr. D. Stone and Dr. G. Gardner for the assistance they gave me with my research. I would also l i k e to thank my wife, Jan, for her continued en-couragement and assistance throughout these sometimes h e c t i c years. 1 INTRODUCTION Much work has been done to demonstrate the ass o c i a t i o n of p a r t i c u l a r species of zooplankton with i d e n t i f i a b l e bodies of water (Russel, 1935, 1936a,b, 1939; Bary, 1959, 1963a,b,c, 1964; Fager and McGowan, 1963). D i f -ferences between bodies of water have been demonstrated i n terms of d i f f e r -ent s u r v i v a l s of organisms i n them (Wilson, 1951). I n i t i a l l y the attempts at d e f i n i n g these differences were r e s t r i c t e d to large scale variables such as s a l i n i t y and temperature. Bary (1963a,b,c) suggested that zooplankton -water body associations were due, instead, to v a r i a t i o n s i n the other pro-p e r t i e s of the water but was unable to define any known chemical or phy s i c a l parameters of the water body which correlated with them. However, changes i n the water q u a l i t y alone have been a t t r i b u t e d to trace amounts of b i o l o g i -c a l l y important compounds (Lucas, 1958; P r o v a s o l i , 1963; Barber and Ryther, 1969; Lewis et a l . , 1971). The b i o l o g i c a l importance of metals i s recognized (Williams, 1953; Bowen, 1966). The term ' b i o l o g i c a l a v a i l a b i l i t y ' of metal implies not only a concentration but also a p a r t i t i o n i n g of the metal which a f f e c t s the a b i l i -ty of the organism to obtain the metal (Lewis et a l . , 1972, 1973). The uptake of metal by marine organisms and e f f e c t s on b i o l o g i c a l systems i s believed dependent upon the form of the metal (Steeman-Nielsen and Wium-Anderson, 1970; Lewis e_t a l . , 1972). The form i s presumably a r e s u l t of physio-chemi-c a l factors r e l a t e d to the metal, the seawater, and the organism (Patin, 1973). I t has been suggested that the predominant form of c e r t a i n metals i n natural waters may be an organic complex (Slowey et a l . , 19,67; Williams, 1968; Slowey and Hood, 1971) although there i s considerable disagreement (Zirino 2 and Healy, 1970; Z l r i n o and Yamamoto, 1972). Johnson (1964) notes that organic material i s responsible f o r maintaining a c e r t a i n l e v e l of metal i n s o l u t i o n thus preventing i t s removal by p r e c i p i t a t i o n . Barber and Ryther (1969); Barber et a l . (1971); Tabata and Nishikawa (1969); Lewis et a l . (1972, 1973) have noted the a b i l i t y of n a t u r a l l y oc-curring organics to reduce the toxi c e f f e c t s of some metals and Lewis et. a l . (1971) showed that the addition of a synthetic chelating agent increased s u r v i v a l of a test organism. At the same time, however, Lewis e_t a l . (1971) noted that a decrease i n s u r v i v a l was af f e c t e d by these same addi-tions at d i f f e r e n t times of the year. I f such a response was an annually repeated phenomenon i t would be reasonable to consider that i t could be link e d with some seasonal occurrence leading to a change i n some b i o l o g i c a l -l y important parameter(s) of water q u a l i t y . (Investigations of factors a f-fe c t i n g primary production show an example of t h i s Barber and Ryther (1969); Barber et a l . (1971)). They found a marked decrease i n s p e c i f i c growth rates concurrent with the appearance of newly upwelled water i n both the Cromwell Current upwelling and the Peru upwelling. I t should be noted that both of these upwellings are induced by seasonal changes i n wind d i r e c t i o n . The present study i s an examination of the r e l a t i o n s h i p between the s u r v i v a l of a bioassay organism and the b i o l o g i c a l a v a i l a b i l i t y of metals; the b i o l o g i c a l a v a i l a b i l i t y of metals and the presence of organics i n the seawater; the presence of v a r i a t i o n i n the organic load of the seawater and seasonal changes i n selected parameters i n the water. In t h i s way i t may be possible to take a further step towards defining a parameter, ch a r a c t e r i s -t i c of a p a r t i c u l a r water type, which may be rela t e d to zooplankton s u r v i v a l . 3 A knowledge of the b i o l o g i c a l properties of Juan de Fuca S t r a i t water i s important to an understanding of the ecology of the S t r a i t of Georgia. Juan de Fuca S t r a i t i s the primary entrance of the S t r a i t of Georgia to the open ocean, the only other entrance being at the north at Johnstone S t r a i t (Waldichuk, 1957). The S t r a i t of Georgia receives deep water of oceanic o r i g i n , and a mixture of t h i s water with i t s own water from Juan de Fuca S t r a i t . The geography of Juan de Fuca S t r a i t i s described by Herlinveaux and T u l l y (1961) as a submarine v a l l e y extending from the ocean (Cape F l a t t e r y ) to the channels of the San Juan Archipelago (Fig. 1). The s t r a i t contains two basins of depths greater than 100 m, separated by a s i l l which l i e s southward from V i c t o r i a , B.C. at 60 m depth. The "Inner S t r a i t " contains several shallow banks, through which the deepest channel leads into Haro S t r a i t and Admiralty I n l e t . The "Outer S t r a i t " deepens gradually to sea-ward to more than 200 m at Cape F l a t t e r y . Beyond the Cape t h i s v a l l e y turns southward at right angles and cuts through the continental slope to the ocean abyss. Deep waters of Juan de Fuca S t r a i t are continuous with ocean waters below 100 m depth (Herlinveaux and T u l l y , 1961), therefore, ocean water moves into the outer part of Juan de Fuca S t r a i t and, being more dense, runs under the upper, low s a l i n i t y , water. Intruding along the bottom t h i s ocean water migrates up the canyon. As the channel of the outer s t r a i t i s s l i g h t -l y curved the 'flood' flow tend to be more on the southern side due to the combination of both the c o r i o l i s and c e n t r i f u g a l forces. The water then moves over the s i l l , through the inner basin, r i s e s up the seaward slope of the ridge and cascades into the S t r a i t of Georgia. The 'ebb' flow i n Juan 4 FIGURE 1. Juan de Fuca S t r a i t and surrounding waters showing the posi t i o n s of JF7-3, GEO 1748 and Cape Beale. 5 125° 124° 123° 6 de Fuca S t r a i t i s concentrated on the northern side of the I n l e t and i s composed of surface waters augmented by runoff waters which vary season-a l l y . vThe largest source of fresh water i n the system i s the Fraser River which provides 70-75% of the fresh water the S t r a i t of Georgia receives annually (Herlinveaux and T u l l y , 1961). Ingelsrud, Robson and Thompson (1936) and T u l l y (1942) have suggested that colder, more s a l i n e water appears at mid-depths i n Juan de Fuca S t r a i t during the summer months. I t i s during t h i s period and the preceding spring months that the p r e v a i l i n g winds change from a S-SE o r i g i n to one of N-NW (Herlineveaux and T u l l y , 1961; T u l l y , 1942; Waldichuk, 1957; Barnes et a l . , 1972; Bakun, 1973) ( F i g . 2). As i l l u s t r a t e d i n Sverdrup et a l . (1942) the N-NW winds blowing along the coast w i l l lead to the trans^ port away from the coast of the l i g h t surface water and, owing to the con-t i n u i t y of the system, t h i s l i g h t water i s replaced by heavier subsurface water. This upwelled water originates on the continental s h e l f . a t .depths of 200-300 m (Doe, 1955). Winds blowing i n the opposite d i r e c t i o n along the coast (S-SE) w i l l lead to an accumulation of l i g h t water along the coast. During the period, October, or November, through to February when S-SE winds are most prevalent and upwelling ceases, a near surface counter cur-rent (the Davidson Current) (Fig. 3) develops r e s u l t i n g i n northerly flow along the coast at a l l depths (Sverdrup e_t a l . , 1942) . This r e s u l t s i n the Northward transport of near surface water from the Columbia River Estuary into Juan de Fuca S t r a i t and even the S t r a i t of Georgia (Barnes e_t a l . , 1972) (Fig. 4). Figure 3 summarizes the major ocean currents prevalent i n the north east P a c i f i c Ocean which may cause v a r i a t i o n i n the composition of water 7 FIGURE 2. Average speed and d i r e c t i o n df monthly winds f or the period 1961-1963 (Barnes et a l . , 1972). 130° 128° 126° 124° 122° 9 FIGURE 3. Shows the major ocean currents of the Northeast P a c i f i c Ocean (Gardner, 1976). 11 FIGURE 4. Location of releases, hypothetical paths, and approximate recovery pos i t i o n s of seabed d r i f t e r s found between the Columbia River and Vancouver Island, more than 100 km from release point (Barnes e_t a l . , 1972) . 13 entering Juan de Fuca S t r a i t . The major e a s t e r l y current i s the West Wind D r i f t which diverges as i t approaches the coast. A small portion of t h i s water may intrude i n t o coastal waters (Dodimead et a l . , 1963) the possible r e s u l t being the movement of near surface water from the Central Subarctic Domain into Juan de Fuca S t r a i t . Water from the south-ern C a l i f o r n i a Undercurrent Domain may also be brought seasonally to the S t r a i t from the subtropics (Dodimead e_t a l . , 1963) . During the upwelling season the northward flow occurs only at depths of 200 m or greater (Sverdrup and Fleming, 1941). With such a number of p o t e n t i a l sources of water for Juan de Fuca S t r a i t consideration must be given to the p o s s i b i l i t y that the character-i s t i c s of the resultant water entering the S t r a i t may vary d r a s t i c a l l y i n t h e i r b i o l o g i c a l impact. The Juan de Fuca S t r a i t IOUBC ( I n s t i t u t e of Oceanography, University of B r i t i s h Columbia) Station 7-3 (48° 17.8'N, 124° 06.3'W) (Fig. 1). was chosen due to i t s p o s i t i o n , s l i g h t l y more than halfway through the S t r a i t . This was intended to provide a 'buffer' distance from s t r i c t l y oceanic conditions and to give water that could l a t e r enter the S t r a i t of Georgia system. Here i t was f e l t the parameters of the Juan de Fuca S t r a i t Deep Water could be examined i n an attempt to c o r r e l a t e bioassay r e s u l t s with the p h y s i c a l and chemical c h a r a c t e r i s t i c s of the seawater. The i n v e s t i g a t i o n of the b i o l o g i c a l a v a i l a b i l i t y of metals was under-taken i n Juan de Fuca S t r a i t and e n t a i l e d : 1. An examination of the b i o l o g i c a l e f f e c t s of metal a v a i l -a b i l i t y on a bioassay organism, the pre-feeding embryo and naupliar stages of Pareuchaeta elongata Esterl e y , using natural seawater from Juan de Fuca S t r a i t . To accom-p l i s h t h i s the seawater was subjected to UV photo-oxidation, to oxidize dissolved matter and l i b e r a t e the metal i n these phases (Williams, 1968). Additions of cop-per, zinc, i r o n and manganese (as chlorides) as w e l l as synthetic chelating agent EDTA (ethylenediaminetetraace-t i c acid) were made to both the natural and UV photo-oxidized seawater to produce conditions of elevated metals and complexing agents. EDTA s u r v i v a l i n enriched, UV photo-oxidized water gives some estimate of the complexing a b i l i t y of the natural organics present i n the seawater. Analysis of Juan de Fuca S t r a i t seawater, both natural and UV photo-oxidized, f o r DOC (dissolved organic carbon) and the added metals, copper, zin c , i r o n and manganese. This to better allow.... Co r r e l a t i o n of the b i o l o g i c a l a v a i l a b i l i t y of metals with the presence of organics i n the seawater. S u p e r f i c i a l examination of environmental factors such as seasonal winds, upwelling, r i v e r runoff, and b i o l o g i c a l pro-d u c t i v i t y which may contribute to changes i n the metal and organic load and subsequent b i o l o g i c a l a v a i l a b i l i t y of metals. 15 MATERIALS AND METHODS i ) In the F i e l d : Water was c o l l e c t e d from hydrographic depths (Table I) at the Juan de Fuca S t r a i t Station, JF7-3 (48° 17.8'N, 124° 06.3'W) (Fig. 1), f o r s a l i n i t y , temperature, and dissolved oxygen measurements. The c o l l e c t i o n was made with NIO b o t t l e s (National I n s t i t u t e of Oceanography, Wormley, UK) equipped with reversing thermometers for temperature determination. S a l i n i t y was determined by measuring conductivity using the Autolab Induc-t i v e l y Coupled Salinometer (Model 301, MK3) and dissolved oxygen by a modified Winkler technique ( C a r r i t t and Carpenter, 1966). Water for DOC (dissolved organic carbon) and metal an a l y s i s , and b i o -assay purposes was c o l l e c t e d from 150 m depth with a 9 6 - l i t e r f i b e r g l a s s and Leucite sampler. Within two hours of c o l l e c t i o n the water was f i l t e r e d with a membrane f i l t e r (0.45 urn, mean pore size) and stored i n acid washed (0.1 N HCl) and seawater rinsed 2 3 - l i t e r polyethylene containers at 8°C u n t i l used. Eggs and ovigerous females of the bioassay organism, Pareuchaeta  elongata E s t e r l e y ( t i t l e page p l a t e ) , were c o l l e c t e d at the S t r a i t of Georgia Station, GS-1748 (49° 17.1'N, 123° 48.0'W) (Fig. 1), with a 1 meter diameter c o n i c a l net (mesh aperture approximately 0.7 mm square) towed v e r t i c a l l y to the surface from approximately 400 m depth. Eggs and ovigerous females were sorted from the rest of the plankton and placed i n thermos f l a s k s , containing GS-1748 water c o l l e c t e d from 350 m depth, for transport back to the laboratory. i i ) Bioassays: The bioassay organism, Pareuchaeta elongata E s t e r l e y (Crustacea; Copepoda; Calanoida) (see appendix) i s widely d i s t r i b u t e d through the North 16 TABLE I. Station l o c a t i o n s , cruise numbers and dates f o r the cruise on which hydrographic data, bioassay organisms and water were c o l l e c t e d . Hydrographic data and water for metal and DOC analyses and bioassay purposes was c o l l e c t e d at Juan de Fuca Station 7-3. A l l bioassay organisms were c o l l e c t e d at S t r a i t of Georgia Station 1748. Hydrographic casts were made to the hydrographic depths given below. STATION ABBREVIATION LATITUDE LONGITUDE Juan de Fuca S t r a i t Station 7-3 JF7-3 48° 17.8'N 124° 06.3'W S t r a i t of Georgia Station 1748 GEO 1748 49° 17.1'N 123° 48.0'W IOUBC CRUISE NO. DATES ABBREVIATION 76-2 February 9-10 (1976) FEB 76-3 March 1-2 » MARC^ 76-5 March 29-30 MARCH2 76-6B May 6-7 MAYX 76-8 May 25-26 MAY 2 76-11 June 14-15 JUNE HYDROGRAPHIC DEPTHS 0 m 10 m 20 m 30 m 50 m 75 m 100 m 125 m 150 m 175 m 200 m 17 P a c i f i c (Davis, 1949). The l i f e h i s t o r y consists of an embryonic phase, 6 naupliar stages (Fig. 5), and 6 copepodite stages (Campbell, 1934). The adult female produces a c l u s t e r of 8-24 deep blue eggs which remains at-tached to the g e n i t a l segment for at l e a s t part of the embryonic develop-ment ( t i t l e page p l a t e ) . The embryo and f i r s t two naupliar (prefeeding) stages u t i l i z e stored reserves f or development whereas the t h i r d through s i x t h nauplius r e l y , to some extent, on phytoplankton (A.S. Pandyan, unpubl.). Previous work by Lewis and Ramnarine (1969) indicates that the embryo and f i r s t two naupliar stages form a c r i t i c a l phase i n the development of P_. elongata. They have since been used i n numerous bioassays, as i n d i c a t o r s of metal t o x i c i t y (Lewis ejt a l . , 1971, 1972, 1973; W h i t f i e l d and Lewis, 1976). On returning to the laboratory from sea 2 egg c l u s t e r s were placed i n 1 - l i t e r polypropylene Erlenmeyer flasks containing 600 ml of seawater. Three f l a s k s were used for each treatment, every f l a s k having f i r s t been rinsed out with 0.1 N HC1 and then with seawater before f i l l i n g . Metal enrichment was with CuC^, ZnC^, TeCl^, or MnC^ dissolved i n deionized, g l a s s - d i s t i l l e d water. A l l stock solutions f o r metal addition were 1000 ppb metal and su i t a b l e aliquots were placed i n 600 ml of seawater to a r r i v e at the required l e v e l s . Additions of the a r t i f i c i a l complexing agent EDTA (ethylenediaminetetraacetic acid) were made from a 500 y.M EDTA stock s o l u t i o n . This stock s o l u t i o n was made by d i s s o l v i n g Na^EDTA i n deionized, g l a s s - d i s t i l l e d water and su i t a b l e aliquots were added to 600 ml of seawater to a r r i v e at the required l e v e l s . The l e v e l s of metals and EDTA added were chosen i n an attempt to give an idea of the e f f e c t s of the addition of a small amount of metal or EDTA vs. a r e l a t i v e l y large amount. 18 FIGURE 5. Naupliar stages of Pareuchaeta elongata E s t e r l e y : a, F i r s t nauplius, dorsal view; b, Second nauplius, dorsal view; c-f, t h i r d through s i x t h n a u p l i i , v e n t r a l views (Lewis and Ramnarine, 1969). 0.5mm 20 U l t r a - v i o l e t i r r a d i a t i o n of seawater was done to photo-oxidize dissolved organic material and c o l l o i d a l matter to l i b e r a t e metals i n -volved i n metal-organic complexes (Beattie et^ a l . , 1961; Armstrong et^ a l . , 1966; St r i c k l a n d , 1972; Williams, 196"8). I r r a d i a t i o n was for a s i x hour period with a 1200 Watt U.V. l i g h t i n a double-walled immersion tube (Hanovia Englehardt Model 189A) immersed i n seawater i n a 5 0 - l i t e r glass chromatographic chamber. The chromatographic chamber was wrapped i n f o i l to reduce l i g h t loss and was covered with plate glass to reduce evapora-t i o n and contamination. The s i x hour treatment was found to reduce the natural complexing a b i l i t y of the seawater organics to a minimum without loss of metals used i n the study (Whitfield, 1974). The water was stored a f t e r UV photo-oxidation, i n acid washed (0.1 N HCl), seawater rinsed 23-l i t e r polyethylene containers at 8°C u n t i l used. The e f f e c t s of various treatments and enrichments of the seawater were measured by comparison of the percent s u r v i v a l to the t h i r d naupliar stage. Six egg cl u s t e r s were used for each treatment i n a monthly s e r i e s . Based on the s t a t i s t i c a l analysis c a r r i e d out by W h i t f i e l d (1974) t h i s num-ber of r e p l i c a t e s allows examination of the e f f e c t s of the treatment of the seawater, while maintaining a standard deviation of les s than 5%. In eva-l u a t i n g the r e s u l t s a difference between means greater than two standard deviations (10%) was considered to be s t a t i s t i c a l l y s i g n i f i c a n t . i i i ) DOC Analysis: Natural (control) seawater and UV photo-oxidized seawater were analyzed f o r dissolved organic carbon (DOC) using the method described by Menzel and Vacarro (1964) with the exceptions noted below: 21 a) 10 ml of sample were used instead of 5 ml, b) glass ampoules were precombusted at 550°C A f t e r r i n s i n g a 50 ml glass syringe and cannula with deionized, glass-d i s t i l l e d water and sample seawater, 50 ml of the sample was drawn into the syringe. A f i l t e r adapter containing a 2.4 cm Gelman Type A, glass f i b e r . f i l t e r (precombusted i n a muffle furnace at 550°C for 4 hours) was placed on the syringe and rinsed with 10-15 ml of the sample water i n the syringe. Aliquots of 10 ml of sample water were then passed through the f i l t e r into three 10 ml glass ampoules which had been precombusted at 550°C fo r four hours (the ampoules contained 0.2 gm of potassium persulphate (K2S2O) and 0.25 ml of 6% phosphoric acid s o l u t i o n ) . c) f i l l e d ampoules were sealed using a s p e c i a l l y fabricated apparatus ("Total Carbon System", Oceanography International Corp. Model 0524B (sealing unit)) with an oxygen-propane flame to prevent CO2 contamination, d) sealed ampoules were heated for 4 hours i n an autoclave at 125°C. DOC was measured, as CO2, with the "Total Carbon System" (Oceanogra-phy International Corp. Model 0524B (Infra-red analysis u n i t ) ) . Blanks to determine reagent and background CO2 were run with a l l batches of sam-ples . iv) Metal Analysis: Seawater samples for copper, zin c , i r o n , and manganese analysis were taken from untreated and UV-treated water i n the 2 3 - l i t e r polyethylene containers, placed i n 1 - l i t e r (copper, zi n c , manganese) or 250 ml (iron) polyethylene b o t t l e s , and a c i d i f i e d to ca. pH 2 with 6N HC1. These were stored i n a r e f r i g e r a t o r at 8°C u n t i l analysed. 22 For analysis the 1 - l i t e r samples were placed i n glass erlenmeyer fl a s k s and the pH was adjusted to pH 7.5 - 8.0 with concentrated ammonium hydroxide (NH^OH). The metals were then extracted into isoamyl acetate with diethyldithiocarbamate on magnetic s t i r r e r s f o r 20 minutes. A f t e r the layers separated, subsequent to s t i r r i n g , the upper layer (isoamyl acetate) was removed. This process was repeated three times. The isoamyl acetate was removed to a 100 ml separatory funnel, the metals being re-extracted i n t o an aqueous phase with chlorine i n 0.1 N HC1. The aqueous phase was c o l l e c t e d into a 25 ml glass stoppered graduated c y l i n d e r and made up to 20 ml. This was then analysed f o r zinc on a Techtron Atomic Absorption Spectrophotometer (no. AA-4). 15 ml of the remaining aqueous so l u t i o n was retained for analysis of copper and manganese. To the 15 ml was added 2 ml of 1.0 M t r i s b uffer (pH ca. 7.0) and 1 ml of a 2% so l u t i o n (by weight) of diethyldithiocarbamate. The metal-diethyldithiocarbamate complex was then extracted i n 5 ml of methylisobutyl ketone (MIBK). This MIBK phase was then analysed on the atomic absorption spectrophotometer for copper and manganese. The amount of absorption obtained for each metal was compared to a standard curve prepared for each metal and a concentra-t i o n f or the o r i g i n a l sample determined ( G r i l l , unpub.). This technique has been shown to have a greater than 95% recovery for a l l metals analyzed ( G r i l l , unpub.). For determination of soluble i r o n the a c i d i f i e d 250 ml samples were treated by the method described by St r i c k l a n d and Parsons (1972). B r i e f l y , 100 ml of sample was reacted with bathophenanthroline i n an acetate buffer i n the presence of hydroxylamine. The coloured ferrous complex thus formed was extracted into isoamyl alcohol and the e x t i n c t i o n of the coloured ex-trac t was measured using a 10 cm c e l l at 5330 £ i n a Perkin-Elmer, Double 23 Beam Spectrophotometer (Coleman Model 124D). v) Wind Data: Wind data for the period June 1975-1976 were obtained from the Department of the Environment, Atmospheric Environment Service, Climato-l o g i c a l Information D i v i s i o n . This data was for the Cape Beale Lighthouse at the western end of Juan de Fuca S t r a i t (Fig. 1) and was i n two forms. The f i r s t was a series of monthly summaries giving the number of hours of wind recorded, by d i r e c t i o n , on a monthly bas i s . These figures were then converted by the author to percents of t o t a l wind recorded per month f or each d i r e c t i o n (N, NE, E, SE, S, SW, W, NW). Data was a v a i l a b l e i n t h i s summarized form for a l l months except November and December 1975 and March to June 1976. The data for these months was i n the form of d a i l y , unsum-marized reports varying i n number from an average of 2.6 reports/day to 6.7 reports/day for any given month (Nov. '75, 2.6/day; Dec. '75, 2.8/day; Mar. '76, 6.3/day; Apr. '76, 6.7/day; May '76, 6.6/day; June '76, 6.2/day). As these reports were generally made on a regular basis ( i n terms of the time of day) the number of reports per d i r e c t i o n was determined and expressed as a percent of the t o t a l number of reports made i n that month. Ekman transport vectors were determined for the period June 1975 -June 1976 from computations of the meridional and zonal components provided by W.P. Wickett of the P a c i f i c B i o l o g i c a l Station, Nanaimo, B.C. (Fisheries and Marine Service). v i ) Fraser River Runoff Data: The discharge of the Fraser River, measured at Hope, B.C. for the period Jan. 1975 - Oct. 1976 was obtained from the Department of the En-vironment, Environmental Management Service, Water Survey of Canada D i v i s i o n . 24 This data was i n the form of d a i l y Discharge i n cubic feet per second. The t o t a l discharge per month was taken and converted to cubic meters 3 3 per second (35.314 f t = 1 m ). v i i ) S t a t i s t i c a l A nalysis: S t a t i s t i c a l analyses of the data were c a r r i e d out using the Univer-s i t y of B r i t i s h Columbia, Biology Data Center, Computer Programmes CCOEF and RPLOG.Program CCOEF calculated c o r r e l a t i o n c o e f f i c i e n t s and r e l a t e d s t a t i s t i c s and RPLOG calculated and plotted simple l i n e a r regressions. 25 RESULTS i ) Bioassays, Hydrography, and DOC and Metal A n a l y s i s : The possible combinations of the bioassay seri e s were used to ex-amine the b i o l o g i c a l a v a i l a b i l i t y of metals and were interpreted i n the following manner: (greater than and les s than imply s i g n i f i c a n t l y ) Condition 1. Survival i n UV photo-oxidized water greater than con t r o l and s u r v i v a l i n metal enriched water greater than c o n t r o l : Metals, e s p e c i a l l y copper, z i n c , i r o n or manganese, are l i m i t i n g due to t h e i r l i m i t e d b i o l o g i c a l a v a i l a b i l i t y . Condition 2. Survival i n UV photo-oxidized water greater than control and s u r v i v a l i n metal enriched water approximates that of the c o n t r o l : Metals, including copper, zin c , i r o n , or manganese, may be l i m i t i n g due to t h e i r l i m i t e d b i o l o g i c a l a v a i l a b i l i t y . Condition 3. Surv i v a l i n UV photo-oxidized seawater greater than control and s u r v i v a l i n metal enriched water le s s than c o n t r o l : Other metals may be low and l i m i t i n g (lower l e v e l s of added copper, zinc, i r o n or manganese may be b e n e f i c i a l ) . UV photo-oxidized organics may be l i m i t i n g by reducing metal a v a i l a -b i l i t y . Condition 4. Surv i v a l i n UV photo-oxidized water l e s s than control and s u r v i v a l i n metal enriched water greater than c o n t r o l : A condition possible i f organic complexing agents are at low l e v e l s and copper, z i n c , i r o n or manganese i s at a l e v e l suf-f i c i e n t l y low to be l i m i t i n g . Other metals are high enough to be l i m i t i n g a f t e r UV photo-oxidation. 26 Condition 5. Surv i v a l i n UV photo-oxidized water less than control and s u r v i v a l i n metal enriched water approximates that of the c o n t r o l : Metals, including copper, zinc, i r o n , or manganese, are not l i m i t i n g or are s l i g h t l y l i m i t i n g due to t h e i r l i m i t e d b i o -l o g i c a l a v a i l a b i l i t y . Condition 6. Survival i n UV photo-oxidized water le s s than control and s u r v i v a l i n metal enriched water l e s s than c o n t r o l : Metals, i n c l u d i n g copper, zinc, i r o n , or manganese, are l i m i t i n g due to excess a v a i l a b i l i t y . Condition 7. Survival i n UV photo-oxidized water approximates that of the control and s u r v i v a l i n metal enriched water greater than that of c o n t r o l : B i o l o g i c a l l y important UV photo-oxidizable organics are at low l e v e l s , copper, zinc, i r o n or manganese a v a i l a b i l i t y i s low, metal concentration i s low and l i m i t i n g . Condition 8. Surv i v a l i n UV photo-oxidized water approximates that of control and s u r v i v a l i n metal enriched water approximates that of c o n t r o l : A condition which completes a l l possible combinations but one d i f f i c u l t to imagine i n the .natural environment. Copper, zin c , i r o n or manganese would not be l i m i t i n g and there would have to be a natural complexing agent which would reduce the toxic e f -fect of the added metals without reducing the a v a i l a b i l i t y of na t u r a l l y occurring copper, zinc, i r o n or manganese. Condition 9. S u r v i v a l i n UV photo-oxidizable water approximates that of the control and s u r v i v a l i n metal enriched water le s s than that i n c o n t r o l : B i o l o g i c a l l y important organics are at low l e v e l s and the b i o l o g i c a l a v a i l a b i l i t y of copper, zin c , i r o n or manganese i s excessive. The a p p l i c a t i o n of these combinations i s shown i n Table I I I . Addition of several concentrations of EDTA to control and UV photo-oxidized water provided supplementary information to that provided i n Table I I I . February; Metals, including copper, zinc, i r o n and manganese, are e i t h e r not l i m i t i n g or only s l i g h t l y l i m i t i n g i n February (Table I I I ) . In comparison to the control there i s a s i g n i f i c a n t decrease i n s u r v i v a l by UV photo-oxidation of the water. Addition of the a r t i f i c i a l complexing agent EDTA seems to combat t h i s decrease and r a i s e the s u r v i v a l i n the UV water (Table I I , F i g . 6). EDTA addition to the control water decreases the sur-v i v a l . Addition of copper has no e f f e c t , zinc appears to be b e n e f i c i a l , and i r o n and manganese additions appear detrimental although none of these changes are s t a t i s t i c a l l y s i g n i f i c a n t (Table I I , F i g . 7). UV photo-oxidation was shown to decrease DOC content and increase the apparent copper and i r o n content i n February (Table IV). Values for zinc i n the control and manganese i n the UV water suggested contamination, and were rejected on the basis of comparison with the other data. The hydro-graphic parameters at 150 m were: s a l i n i t y , 33.777 ppt; temperature, 7.60°C oxygen, 2.78/ml/l (Table V, F i g . 8). March^: The hydrographic data (Fig. 8) i s i n d i c a t i v e of a very marked change i n background conditions as the s a l i n i t y decreased (33.349 ppt), tempera-ture decreased (7.38°C), and oxygen increased (3.33 ml/1) (Table V). Changi TABLE II. Percent Survival of the Pre-feeding Stages of P. elongata Eaterley In aeavater collected froo 150 ta depth at' Juan de Fuca Station 7-3 (48° 17.8'N, 124° 06.4*W). Month Control 6 Br. OV 0.5 Copper (lig/1) 1.0 3.0 5.0 1.0 Zinc (wg/1) 5.0 10.0 1.0 Iron (Hg/1> 10.0 15.0 1.0 Manganese (VS/D 10.0 15.0 EDTA (vg/l) 0.02 0.05 0.10 6 0.02 Hr. UV +ED7A (ug/l> 0.04 0.05 0.06 0.10 1976 February 75.9 48.8 71.8 77.2 79.2 81.3 73.2 67.5 74.0 70.5 61.8 68.6 63.5 62.7 48.8 _ 65.4 Marchx 58.8 67.1 63.4 68.4 72.9 74.4 76.5 69.6 61.4 72.4 63.4 72.4 50.6 69.5 71.3 51.7 63.9 64.4 Karchj 58.9 56.1 53.6 59.2 68.7 56.8 66.7 63.7 62.1 45.7 54.7 46.2 74.4 82.0 71.6 46.9 45.6 ,, 54.8 ^ 1 37.7 25.5 46.4 39.6 38.2 43.8 49.5 42.3 58.1 56.4 43.0 59.6 44.9 54.6 53.4 41.1 35.0 L, . 32.0 Kay2 45.2 56.1 51.4 54.1 58.5 55.0 51.8 59.0 53.4 57.0 51.1 38.7 37.1 34.2 30.8 45.3 43.8 . 36.4 June 50.0 33.3 54.2 24.8 42.3 56.8 65.2 46.4 45.6 34.4 48.4 : 55.1 19.8 27.9 45.7 29.9 18.0 34.7 29 TABLE I I I . Combinations of the bioassay series used to examine the b i o l o g i c a l a v a i l a b i l i t y of metals i n Juan de Fuca S t r a i t . Greater than and less than mean that the test r e s u l t s are more than two standard deviations away from those of the control. Combination Copper Zinc Iron Manganese Interpretation of Combination Survival UV > Control and Survival copper, zinc, Iron or manganese > con t r o l Survival UV > Control and Survival copper, zinc, i r o n or manganeses Control 3. Survival UV > Control and Survival copper, zinc, i r o n or manganese < Control 4. Survival UV < Control and Survival copper, zinc, i r o n or manganese > Control 5. Survival UV < Control and Survival coppar, zinc, i r o n or manganese*^ Control 6. Survival UV < Control and Survival copper, zinc, i r o n , or manganese < Control Survival U V ^ Control and Survival copper, zinc, i r o n or manganese > Control 8.. Survival UV«VControl and Survival copper, zinc, Iron and manganese*vcontrol Survival UV«** Control and Survival copper, zinc, i r o n and manganese < Control 2,5 2,5 2,5 2 4,6 4 4 1.4 1 1 1,6 Metals, e s p e c i a l l y copper, z i n c , i r o n or manganese, are l i m i t i n g due to t h e i r l i m i -ted b i o l o g i c a l a v a i l a b i l i t y Metals, including copper, z i n c , i r o n or manganese , may be l i m i t i n g due to t h e i r l i m i t e d b i o l o g i c a l a v a i l a b i l i t y . Other metals may be low and l i m i t i n g (low-er l e v e l s of added copper, z i n c , i r o n or manganese may be b e n e f i c i a l ) . UV photo-oxidized organics may be l i m i t i n g . Copper, zinc, iron or manganese i s at a low l e v e l and other complexed metals are high enough to be l i m i t i n g a f t e r UV photo-oxidation. Metals, including copper, z i n c , i r o n or manganese are not l i m i t i n g or are s l i g h t l y l i m i t i n g due to t h e i r l i m i t e d b i o l o g i c a l a v a i l a b i l i t y . Metals, including copper, z i n c , i r o n or manganese, are l i m i t i n g due to excess a v a i l a b i l i t y . B i o l o g i c a l important UV photo-oxidizable organics at low l e v e l s , copper, zinc, i r o n or manganese a v a i l a b i l i t y low, Metal l i m i t -ing. Copper, zinc, iron or manganese not l i m i t i n natural organics reducing the toxic e f f e c t of the added copper, z i n c , i r o n or mangancs without reducing the a v a i l a b i l i t y of natura l y occurring copper, z i n c , i r o n or manganes B i o l o g i c a l l y important organics are at low l e v e l s , b l o l o g i c . i l a v a i l a b i l i t y of copper, zinc, iron or manganese excessive. February (1), March^ (2), Mnrch 2 (3), Mayx (4), May2 (5), June (6) 30 FIGURE 6. Graphs showing s u r v i v a l s of EDTA addition s e r i e s i n control and UV photo-oxidized seawater f or Feb.-June, 1976. 1 0 0 EDTA additions to Control 31 < 5 0 > 10 0 A ?,100 T EDTA additions to 6hr UV  w 1 Control LU CL 5 0 0 L Feb Mar Apr May Jun EDTA Control • . 0 2 J J M / I EDTA * . 0 5 J J M / I Control • .10JJM/I TABLE IV. Background l e v e l s of DOC (Dissolved Organic Carbon) and Metals (Copper, Zinc, Iron and Manganese) i n Seawater c o l l e c t e d from '. 150 m depth at Juan de Fuca Station 7-3 (48° 17 .8'n, 124° 06.4'W). Levels were determined i n na t u r a l and UV photo-oxidized seawater. Analysed Water Type February March^ Marcti^ May^ May 2 June DOC, (mg C/l) Control 6 Hour UV 0.980 0.925 1.165 0.880 0.660 0.565 0.710 0.535 1.150 0.540 0.870 0.470 Copper Control 1.4 2.2 2.0 1.5 2.7 3.8 (yg / D 6 Hour UV 1.8 2.9 1.2 5.7 3.5 2.8 Z i n W i ) Control 60.0+ 6.4 4.8 4.2 ' 5.6 4.7 6 Hour UV 12.2 6.4 7.2 7.3 12.3 5.2 Iron Control 2.8 11.3 9.2 5.3 3.7 6.2 (yg / D 6 Hour UV 9.7 9.0 15.2 9.6 2.4 5.3 Manganese Control 0.9 36.3+ 0.9 1.0 0.6 0.8 (yg / D 6 Hour UV 16.3 37.0+ 1.2 1.5 0.6 1.0 + . data rejected by author on basis of comparison with other data. O J 33. TABLE V. S a l i n i t y , temperature, and oxygen values of the bioassay control water obtained from 150 m at JF7-3 S a l i n i t y (Ppt) Temperature ( C) Oxygen (ml/1.) Feb. Marchj March,. May 2 June 33.777 33.349 33.688 33.871 33.884 33.859 7.60 7.38 6.94 6.72 6.64 6.63 2.78 3.33 2.60 2.50 2.48 2.40 34. FIGURE 7. Graphs showing s u r v i v a l s of metal addition s e r i e s i n control water for Feb. - June, 1976. 1 0 0 -5 0 f 0 : 1001 < > 501 > on 1 0 o H 100 z 111 U cn i i i CL 5 0 0 100 5 0 0 f 4f Copper ® Q5ug/I o 1.0ug/l A 3.0 pg/l . A 50 j jg / l Z inc © 1.0 pg/l o 5.0 pg/l AlQO(jg/ l Iron © 1.0ug/l o lQOj jg / l &150ug/ l Manganese @ 1.0ug/l o10.0ug/l &15.0 ug/ l Feb Mar Ap r May June 36. FIGURE 8. S a l i n i t y , temperature and oxygen f o r Feb - June, 1976 at JF7-3 (48° 17.8'N, 124° 06.3'W). Hatched l i n e at 150 m indicates depth at which water, f o r bioassay and DOC and metal analysis purposes, was taken. (Vi) HidlQ 38. i s also found i n the values of DOC and metals, both of which appear to increase s u b s t a n t i a l l y (Table IV). A l l the metals considered were l i m i t i n g due to t h e i r l i m i t e d b i o -l o g i c a l a v a i l a b i l i t y (Table I I I ) . A sharp decrease i n the control sur-v i v a l i s noticed, combined with an increase i n UV s u r v i v a l to a point higher than that of the control (Table I I ) . There i s a s i g n i f i c a n t i n -crease i n s u r v i v a l over the c o n t r o l induced by a l l the metals (Table I I , Fig. 7). Additions of EDTA to the control yielded a s i g n f i c a n t increase i n s u r v i v a l while s i m i l a r additions to the UV water caused a reduction i n s u r v i v a l , though not s i g n i f i c a n t (Table I I , F i g . 6). March 2: There i s a r e v e r s a l of the hydrographic change exhibited i n March^ data (Fig. 8) the s a l i n i t y increasing and oxygen decreasing (Table V). A decrease i n the metal and DOC values of the control waters i s also noticed (Table IV). Copper l i m i t a t i o n of s u r v i v a l and a possible excess of i r o n are i n d i -cated for March 2 (Table I I I ) . In t h i s month and subsequent months there appear to be s u b s t a n t i a l differences i n the control metal compared to UV water metal content for various metals (Table IV). The c o n t r o l and UV sur-v i v a l s are v i r t u a l l y the same i n Marct^ (Table I I ) , however, the control s u r v i v a l i s increased by the addition of EDTA while the UV s u r v i v a l i s decreased (Table I I , F i g . 6). This i s compatible with the low l e v e l s of UV photo-oxidizable organics experienced (Table IV). Additions of copper appeared marginally b e n e f i c i a l , zinc caused no s i g n i f i c a n t change, ir o n and manganese both caused a decrease i n s u r v i v a l (Table I I , F i g . 7). 39. May_1: Low and l i m i t i n g l e v e l s of zinc, i r o n and manganese were experienced i n May^ while other metals, i n c l u d i n g copper, are not l i m i t i n g or are only s l i g h t l y l i m i t i n g (Table I I I , Table I I , F i g . 7). A f t e r UV photo-oxidation some metal l e v e l s are a v a i l a b l e enough to be s l i g h t l y i n excess (Table I I I , Table I I ) . This fact i s further i l l u s t r a t e d by the EDTA addition s e r i e s (Table I I , F i g . 6) where additions to both water types r e s u l t s i n an i n -creased s u r v i v a l . Both the control and UV s u r v i v a l s are sharply reduced r e l a t i v e to Marct^ (Table I I ) . Metal additions resulted i n increased sur-v i v a l (although the copper induced increase was not s i g n i f i c a n t , i t was marginal) (Table I I , F i g . 7). It i s during t h i s period that the i n t r u s i o n of deep, more s a l i n e , low-er temperature and oxygen water becomes more apparent (Fig. 8, Table V). It i s marked by a decrease i n metal concentrations, with the exception of manganese, and an increase i n DOC (Table IV). May ^ • The deep wedge of intruding high s a l i n i t y , low temperature and oxygen water i s well established (Fig. 8). The background DOC, copper and zinc values have increased while the manganese and i r o n values have dropped (Table IV). As i n March^, May^ i s a period i n which metals are low and l i m i t i n g and DOC l e v e l s are markedly higher (Table I I I , Table IV, F i g . 7). UV photo-oxidation increased the s u r v i v a l above that of the control (Table II) while EDTA additions to both UV water and control water appear d e t r i -mental (Table I I , F i g . 6). Additions of copper, zin c , and i r o n increase s u r v i v a l (Table I I , F i g . 7). 40 June: This period i s marked by the beginning of the recession of the intruding high s a l i n i t y , low oxygen and temperature water (Fig. 8, Table V). This i s p a r a l l e l e d by lower DOC and zinc values and higher amounts of copper, i r o n and manganese (Table IV). The control s u r v i v a l continues to increase r e l a t i v e to May^ and May^ while the UV s u r v i v a l drops dras-t i c a l l y (Table I I ) . June i s a month when metals are apparently i n excess as indicated by the bioassay r e s u l t s with added copper or i r o n (Table I I I , Table I I , F i g . 7). UV photo-oxidation leads to greatly decreased s u r v i v a l as did EDTA additions to both c o n t r o l and UV water (Table I I , F i g . 6). Additions of copper proved detrimental while zinc was b e n e f i c i a l at low l e v e l s , i r o n decreased s u r v i v a l and manganese had no s i g n i f i c a n t e f f e c t (Table I I , F i g . 7). i i ) Wind Wind data for the period June 1975 - June 1976 have been summarized (Table VI) to give an o v e r a l l p i c ture of the winds which were prevalent during the time of study. Although the data do not provide as d r a s t i c an example of the s h i f t i n wind d i r e c t i o n that occurs i n the spring - summer months as that shown i n Figure 2, i f one considers a summation of the per-cent of the winds which were from a N or NW o r i g i n the increase i s very marked. This s h i f t i s evident i n Figure 9 i n the wind rosettes prepared for t h i s period. Figure 10 shows the calculated Ekman transport values for the area about the mouth of Juan de Fuca S t r a i t . These c l e a r l y show the s h i f t i n water transport which occurs i n the l a t e spring and summer months. TABLE VI. Gives the percent of the wind which blew i n each d i r e c t i o n per month f o r the period June 1975 - June 1976. The under-l i n e d percentage i n each month indicates the d i r e c t i o n of the p r e v a i l i n g wind for that month. MONTH DIRECTION TOTAL N NE E SE S SW W NW CALM HOURS/ (percent of t o t a l hours reported) REPORTS June 1975 1.9 2.4 1.0 26.0 11.8 8.6 33.2 14.9 0.3 720* July II 1.9 0.7 1.6 28.8 10.3 5-9 31.9 15.9 3.1 744* August II 0.8 2.0 1.6 42.6 10.6 5.1 24.3 9.8 3.1 744* September II 10.39 8.7 6.7 28.5 11.9 6.0 14.3 7.3 6.0 712* October II 1.5 8.2 7.9 48.4 9.5 10.2 9.4 4.3 0.5 744* November II 0.0 25.6 5.1 38.5 7.7 10.3 9.0 3.8 0.0 78+ (2.6) December II 4.7 17.4 11.6 29.1 3.5 10.5 8.1 8.1 7.0 86+ (2.8) January 1976 5.2 10.3 11,7 34.1 6.9 10.7 12.4 7.5 1.1 735 February II 1.6 15.9 10.3 26.1 8.6 14.2 17.5 4.7 0.9 696* March II 1.5 15.8 4.6 29.6 8.7 9.2 14.8 11.2 4.6 .196+ (6.3) A p r i l II 2.5 13.9 4.5 34.3 2.5 6.0 13.9 15.4 7.0 201+ (6.7) May II 1.5 8.7 1.9 31.6 3.9 9.2 24.3 14.6 4.4 206+ (6.6) June II 2.7 2.7 2.7 20.9 10.7 9.1 29.4 16.6 5.3 187+ (6.2) report data taken from summarized monthly reports. Data i n form of hours of wind per d i r e c t i o n , converted here to a percent of t o t a l hours. The t o t a l num-ber of hours reported for each month i s indicated at r i g h t . report data taken from unsummarized d a i l y reports. Number i n far r i g h t column i n t h i s case indicates the t o t a l number of reports made during the month. This number was compared with the number of reports f or that month i n any given d i -r e c t i o n to a r r i v e at the percent value for that d i r e c t i o n . Numbers i n brackets indic a t e the average number of reports per day for that month. 42. FIGURE 9. Gives the wind d i r e c t i o n s (percent of monthly t o t a l ) experienced at Cape Beale f o r the period June 1975 - June 1976. 43. y% 3 w 17^5 5 yY £) i , n J V n e 30(-rS)\ June s 4 8 . 0 5 % W + N W 4 5 . 9 8 % 15,3 J u l y ' 7 5 W+NW 40 4 7 7 1 o / o 29 5 A u g ' 7 5 • ' | 0 W+NW 'J® 3 4 . 1 3 % 50-11 9 / ^ ° S e p ' 7 5 QV$feo W + N W '28 2 1 . 6 2 % 5X8 Oct ' 75 1 0(tj9| 0 W + N W 1 1 N p V 0 13 .70% 1 0 < ® % W + N W 1 6 . 2 6 % 16V3 ;3 M a y ' 7 6 20 W t N W y$ 3 8 . 8 3 % 1 7 < « 5 A p r ' 7 6 15\iS5o W + N W ^£XJ° 2 9 . 3 5 % 12/^2S]5 1 6 f t § ? $ . ^ r ' 7 6 2>29 26.01°/ . 9 V^Lxio W + N W i T V Q ^ g 0 2 2 . 2 6 % l 2 / 5 v ^ 1 2 J a n ' 7 6 W + N W 1 9 . 8 6 % ^°^>33 44. FIGURE 10. Gives the calculated Ekman Transport values for the 5° grids about 4'5°N, 125°W and 50°N, 130°W f o r June 1975 - June 1976. 0 45. 46. i i i ) Fraser River Runoff Data representative of the discharge from the Fraser River at Hope, B.C., have been depicted i n Figure 11 f o r the period January 1975 to 4 3 -1 October 1976. A peak discharge of 20.0 - 25.0 (X 10 m • sec ) can be seen annually i n the summer months with the lowest discharges of 2.0 - 3.0 (X 10 m • sec ) occurring i n the winter and early spring. There i s a very rapid increase i n outflow between March and May. iv) S t a t i s t i c a l Analysis The c o r r e l a t i o n c o e f f i c i e n t s ( r ) , obtained by c o r r e l a t i o n analysis of the v a r i a b l e s , were calculated and shown i n Table VII. As only a very l i m i -ted number of the c o e f f i c i e n t s were s i g n i f i c a n t at l e v e l s of 1% to 5% (Snedecor, 1956, Table 7.6.1, pp. 174) values of ' t ' were calculated using equation (1) (Snedecor, 1956): t = r- [ ( n - 2 ) / ( l - r 2 ) ] 1 / 2 (1) where: r = c o r r e l a t i o n c o e f f i c i e n t (from Table V) n = number of samples df (degrees of freedom) = n - 2 A 't-Table' (Snedecor, 1956, Table 2.7.1, p. 46) was used to determine the p r o b a b i l i t y of r e j e c t i o n of the n u l l hypothesis (H Q) with respect to the c o r r e l a t i o n of the variables (Table VI I I ) . A high c o r r e l a t i o n was noted i n the data between s a l i n i t y and oxygen, temperature and oxygen ( s i g n i f i c a n c e at 0.5% l e v e l ) ; s a l i n i t y and tempera-ture ( s i g n i f i c a n c e at 1.0% l e v e l ) ; s a l i n i t y and control i r o n , oxygen and control DOC ( s i g n i f i c a n c e at 5.0% l e v e l ) (Table VI I I ) . Of lower s i g n i f i c a n c e 47 FIGURE 11. Shows the monthly Fraser River discharge at Hope, B.C. i n cubic meters per second (X 10^). 48. TABLE VII. Gives the correlation coefficients (r) obtained by correlation analysis of variables using the UBC Biological Computing Center Programme CCOEF. PSCT SAL TEM OXY. SAL -0.382 TEM ; +0.664 -0.905* OXY +0.470 -0.947+ +0.954 CUCT -0.316 +0.183 -0.375 -0.333 ZNCT +0.556 -0.740 +0.710 +0.810 FECT . +0.003 -0.826* +0.555 +0.608 MNCT +0.174 -0.515 +0.507 +0.368 DCCT +0.550 +0.729 -0.792 -0.816* -0.715 +0.301 +0.327 -0.363 -0.229 +0.669 +0.686 CUCT ZNCT FECT MNCT -0.088 +0.613 +0.394 +0.289 -0.581 +0.248 +0.110 -0.658 +0.705 -0.352 +0.320 +0.229 +0.121 PSUV CUUV ZNUV FEUV MNUV SAL TEM' OXY CUUV ZNUV FEUV MNUV DCUV Note: key to variable name abbreviations i n Table IX. * correlation coefficient significant at the 5% level. + correlation coefficient significant at the 1% level. TABLE V T I I . Gives the p r o b a b i l i t y of rej e c t i o n of the null-hypothesis (H q), with respect to the c o r r e l a t i o n of the variables, as derived from a table of the d i s t r i b u t i o n of ' t ' (Snedecor 1956). Values of ' t ' were calculated from the correlation c o e f f i c i e n t s (r) obtained by corr e l a t i o n analysis of variables using the UBC B i o l o g i c a l Computing Center Programme CCOEF. PSCT SAL TEM • OXY CUCT ZNCT FECT MNCT SAL 0.500 TEM . 0.200 0.010 OXY 0.400 0.005 0.005 CUCT >0.500 >0.500 0.500 >0.500 ZNCT 0.400 0.200 0.200 0.100 -• FECT >0.500 0.050 0.400 0.200 M^ CT >0.500 0.400 0.500 >0.500 DCCT 0.200 0.100 0.100 0.050 >0.500 0.400 0.500 >0.500 0.150 >0.500 >0.500 0.500 >0.500 0.200 0.150 0.400 >0.500 >0.500 0.400 0.200 0.500 > 0.500 > 0.500 > 0.500 PSUV CUUV ZNUV FEUV MNUV SAL-. TEM : OXYV CUUV ZNUV FEUV MNUV DCUV Note: key to variable name abbreviations i n Table IX. o 51 were cor r e l a t i o n s between s a l i n i t y and co n t r o l DOC, temperature and con-t r o l DOC ( s i g n i f i c a n c e at 10.0% l e v e l ) ; s a l i n i t y and s u r v i v a l i n the UV control bioassay, oxygen and s u r v i v a l i n the UV c o n t r o l bioassay ( s i g n i -ficance at 15.0% l e v e l ) ; s u r v i v a l i n co n t r o l bioassay and temperature, s u r v i v a l i n control bioassay and control DOC, s a l i n i t y and co n t r o l z i n c , temperature and co n t r o l zinc, oxygen and co n t r o l i r o n , temperature and s u r v i v a l i n UV c o n t r o l , UV co n t r o l DOC and s u r v i v a l i n the UV control ( s i g n i f i c a n c e at 20.0% l e v e l ) (Table V I I I ) . The sign, p o s i t i v e (+) or negative (-), of the c o r r e l a t i o n c o e f f i -cients (r) given i n Table VI i s i n d i c a t i v e of the type of r e l a t i o n s h i p e x i s t i n g between the v a r i a b l e s . A summary of s i g n i f i c a n t c o r r e l a t i o n s has been made i n Table X showing the r e l a t i o n s h i p s e x i s t i n g between the v a r i -ables. 52 TABLE IX. Provides a key to the v a r i a b l e name abbreviations used i n the s t a t i s t i c a l analysis of the r e s u l t s . ABBREVIATION PSCT percent s u r v i v a l of the control PSUV percent s u r v i v a l of the UV control SAL s a l i n i t y values f o r the period February 1976-June 1976 TEM temperature values f or the period February 1976-June 1976 OXY oxygen values f or the period February 1976-June 1976 CUCT copper content of the control water CUUV copper content of the UV photo-oxidized water ZNCT zinc content of the control water ZNUV zinc content of the UV photo-oxidized water FECT i r o n content of the control water FEUV ir o n content of the UV photo-oxidized water MNCT manganese content of the control water MNUV manganese content of the UV photo-oxidized water DCCT DOC content of the cont r o l water DCUV DOC content of the UV photo-oxidized water 53 TABLE X. A summary of the s i g n i f i c a n t correlations. The sign placed in the table indicates whether the relationship between the two variables i s pos i t i v e or negative. PSCT PSUV SAL TEM OXY ZNCT FECT DCCT DCUV PSCT PSUV SAL -TEM + + -OXY . + - + ZNCT - + FECT - • ' + DCCT + + - ' DCUV + Table IX - key to variable name abbreviations 54 DISCUSSION Changes i n water q u a l i t y have been r e l a t e d to subsequent, changes i n the growth or s u r v i v a l of organisms resident i n that water. Two periods of abrupt change i n water character are indi c a t e d i n the hydrographic data (Fig. 8). One during February - A p r i l and one during May - June. Water f o r the March^ bioassay came from the f i r s t , water for the May 2 b i o -assay came from the second. The March^ change appears to have res u l t e d i n a mixing of the e n t i r e water column whereas the May 2 water character-i s t i c s appear to r e s u l t from an i n f l u x of upwelled water of deep, oceanic o r i g i n ( F ig. 8). It i s suggested that colder, more s a l i n e water appears at mid-depths i n Juan de Fuca S t r a i t during the summer months (Ingelsrud, Robson and Thompson, 1936; T u l l y , 1942). I t i s during t h i s period, and the preceding spring months, that the p r e v a i l i n g winds change from a S-SE o r i g i n to one of N-NW (Herlinveaux and T u l l y , 1961; T u l l y , 1942,5 Waldichuk, 1957; Barnes et a l . , 1972; Bakun, 1973) ( F i g . 2). An increase i n W-NW winds i s also apparent i n the spring of 1976 (Fig. 9). T u l l y (1942) c a l -culates that during t h i s period the steady state of the wind driven cur-rents necessary for upwelling water at the mouth of Juan de Fuca S t r a i t should be attained i n les s than a day. Since there are from four to ten major changes of wind per month, each major wind has time to e s t a b l i s h a steady current. W-NW winds that produce upwelling being subsurface water from depths of 200 m - 300 m from the continental s h e l f (Doe, 1955). Bakun (1973) has calculated an "upwelling index" based on the mathematics of Ekman Transport using data for the twenty year period, 1948-1967 (Fig. 12a, b). These indi c a t e the p o s s i b i l i t y of upwelling occurring i n the 55 spring and summer months at the mouth of Juan de Fuca S t r a i t . Consulting F i g . 10 one finds the s h i f t i n Ekman Transport, leading to upwelling, to be i n evidence i n the l a t e spring and summer of 1976. The peak outflow of the Fraser River occurs during the l a t e spring -summer upwelling period (Fig. 11), increasing entrainment of deep Juan de Fuca S t r a i t water. This entrainment by i t s e l f i s not s u f f i c i e n t to cause the d r a s t i c change i n the hydrographic character noted within the S t r a i t but i t w i l l f a c i l i t a t e the upwelling and seaward transport of any water i n Juan de Fuca S t r a i t . S t a t i s t i c a l analysis of the data suggests a r e l a t i o n s h i p between the hydrographic c h a r a c t e r i s t i c s and the presence of DOC. Table X shows a p o s i t i v e c o r r e l a t i o n between DOC i n the bioassay control water and the sa-l i n i t y of the water (p = 0.100) and a negative c o r r e l a t i o n between DOC and the temperature and oxygen content of the same water (p = 0.100 and p = 0. This c o r r e l a t i o n suggests that the higher s a l i n i t y , colder, upwelled water would have a higher organic content. The analysis further shows the DOC of the control water i s negatively correlated with the s u r v i v a l of the control bioassay (p = 0.200) (Table X). The s u r v i v a l of the UV control bioassay i s also correlated with the DOC of the UV photo-oxidized water (p = 0.200) and s a l i n i t y (p = 0.150), temperature (p = 0.200) and oxygen (p = 0.150) (Table X). There i s also l i m i t e d c o r r e l a t i o n between zinc i n the control and s a l i n i t y (p = 0.200) and temperature (p = 0.200) and between i r o n i n the control and s a l i n i t y (p = 0.050) and oxygen (p = 0.200) (Table X). The b i o l o g i c a l a v a i l a b i l i t y of metals i s c o n t r o l l e d by the amount of the metal that i s present as we l l as the state i n which the metal occurs. 56 FIGURE 12a. Mean monthly values of the "upwelling Index" f o r the 20 year period, 1948-1967. Units are cubic meters per second per 100 m of co a s t l i n e . A p o s i t i v e value i s i n d i c a t i v e of the existence of p o t e n t i a l upwelling conditions. (Bakun, 1973). FIGURE 12b. Shows a graphic demonstration of the "upwelling i n -dex" (above, F i g . 12a). Mean monthly values of the computed upwelling indices for the 20 year period 1948-1967. (Bakun, 1973). ^ S T A T I O N 48°N 125° W J a Fe Ma Ap My J n JI Au Se Oc No 90 -47 -21 0 18 25 3 4 22 4 - 3 9 - 8 8 - 1 0 0 •200 J a I F<21 MalAp lMy lJn U l lAu lSe lOc! Nol De -200 58 With high l e v e l s of organic complexing agents the metal may occur i n a buffered/complexed state (Provasoli, 1963), i t s a v a i l a b i l i t y being regu-la t e d by the equilibrium between the metal and the organics (Lewis, unpub.)- With low l e v e l s of organic complexing agents the soluble metal occurs i n an i o n i c state or i n association with inorganic ligands. In t h i s case the b i o l o g i c a l a v a i l a b i l i t y may be c o n t r o l l e d by the concentration of the metal(s). The l e v e l s and states of n a t u r a l l y occurring metals and organics ap-proaches an optimum during February when background DOC l e v e l s are moder-ate and metals occur i n r e l a t i v e l y low amounts (Table IV). As evidenced by the bioassays metals were e i t h e r not l i m i t i n g or only s l i g h t l y l i m i t i n g during t h i s time (Table I I I , Table IV, F i g . 6). Low b i o l o g i c a l a v a i l a b i l i t y of metal occurred concurrent with markedly higher DOC l e v e l s i n March^ and May 2, the two periods of abrupt change i n hydrographic character (Table I I I , Table IV, F i g . 8). The r e l a t i o n s h i p be-tween low a v a i l a b i l i t y of metals and the presence of high DOC suggests that a component of the DOC may have been responsible for l i m i t i n g the b i o l o g i c a l a v a i l a b i l i t y of metals. The change i n hydrographic properties at March^ suggests that t h i s may have been due to a change i n water q u a l i t y , namely, higher DOC associated with down mixing of nearsurface water. The d i f f e r e n t response to additions of EDTA experienced i n these two periods could be a t -tributed to differences i n the nature of the organic complexing agent(s) present i n the water at these two times of the year. The increase i n sur-v i v a l with EDTA addition i n March^ (Table I I , F i g . 6) could r e s u l t from the EDTA chelation of metal(s) i n a form more sui t a b l e to the organism: 59 i . e . , METAL ORGANIC + EDTA METAL EDTA + ORGANIC Where: ORGANIC = natural organic complexing agent. This would be p l a u s i b l e when the s t a b i l i t y constant f o r the EDTA complex i s stronger than that of the natural organics. In addition to a change i n the state of the metal by EDTA addition, p r e f e r e n t i a l complexation of metals by the natural organics and EDTA (Goldberg, 1957; Schubert, 1954; Johnspn 1964) plus competition for metals between natural and added organics may r e s u l t i n new r a t i o s of b i o l o g i c a l l y a v a i l a b l e metals. For example, Lewis (unpubl.) has shown that a r a t i o of 7:1, copper:zinc, r e s u l t s i n the highest s u r v i v a l at any l e v e l of copper. This suggests a p o t e n t i a l l y complex arrangement between s u r v i v a l and the nature of the metal - organic and metal- metal r e l a t i o n s h i p s i n a body of water. Low l e v e l s of UV photo-oxidizable organics and high l e v e l s of Iron ( r e l -a tive to DOC) experienced i n Marcl^ (Table IV) allowed copper l i m i t a t i o n and an excess of i r o n to a f f e c t s u r v i v a l (Table I I I ) . This i s evidenced by the marked increase i n s u r v i v a l caused by addition of EDTA to the control (Table I I , F i g . 6). Marginal conditions e x i s t i n May^ and June when a balance between metal excess and l i m i t a t i o n appears p a r t l y c o n t r o l l e d by the photo-oxidizable or-ganics present and a number of metals (Table I I I ) . Here, as i n March^ and May 2, the r a t i o s and states of the metals could be of great importance. Pre-f e r e n t i a l complexation of s p e c i f i c metals may occur and any change r e s u l t i n g from addition of metals or photo-oxidation of organics may r e s u l t i n b i o l o -g i c a l l y important changes i n metal r a t i o s and/or states. 60 Changes i n the growth of phytoplankton organisms due to f l u c t u a t i o n s i n water properties have been documented by Barber and Ryther (1969) and Barber et a l . (19 71). Working i n the Peru and Cromwell upwelling currents they noted low s p e c i f i c growth rates of phytoplankton i n nutrient r i c h , newly upwelled water. On the basis of the data assembled here i t i s sug-gested that the lower growth rate noted by Barber and Ryther (1969) and Barber ejt a l . (1971) may have been due to the presence of some organic(s) i n the upwelled water which reduces the b i o l o g i c a l a v a i l a b i l i t y of metals. As the water aged at the surface a f t e r the i n i t i a l upwelling period Barber and Ryther (1969) and Barber e_t a l . (1971) noted a gradual increase i n the rate of primary production due to a change i n water c h a r a c t e r i s t i c s . The gradual increase i n the May to June bioassay co n t r o l s u r v i v a l s , a f t e r the i n i t i a l appearance of upwelled water and subsequent ageing and mixing with near surface waters i n the S t r a i t , may be due to s i m i l a r processes. This could be due to the change of the metal into a more sui t a b l e state. The change i n state may r e s u l t from breakdown of the complexing agent(s) or r e s u l t from a change i n the state of the metal caused by near surface b i o l o g i c a l a c t i v i t y (or both). The seasonal occurrence of increased DOC i n Juan de Fuca S t r a i t can be d i r e c t l y r e l a t e d to the appearance of higher s a l i n i t y , lower temperature and oxygen, upwelled water and increased surface p r o d u c t i v i t y i n the S t r a i t . The mechanism for upwelling being the W-NW winds which predominate the l a t e Spring-Summer period annually. A r e l a t i o n s h i p between the b i o l o g i c a l a v a i l -a b i l i t y of metal i n Juan de Fuca S t r a i t and the presence of organic complex-ing agents has been suggested. In oceanic water, where the natural l e v e l s of metal are low, the introduction of upwelled water, containing or4. g a n i c a l l y complexed metal, reduces the b i o l o g i c a l a v a i l a b i l i t y of metals. 61 This may decrease the s u r v i v a l of organisms resident i n that water. Sub-sequent 'ageing' of the water may lead to a gradual increase i n the sur-v i v a l , possibly through photo-oxidation of the organic ligand(s) and pro-duction of b i o l o g i c a l l y more suitable metal states. 62 BIBLIOGRAPHY Armstrong, F.A.J., P.M. Williams, and J.D.H. Str i c k l a n d . 1966. Photo-oxidation of organic material i n seawater by u l t r a - v i o l e t r a d i a t i o n , a n a l y t i c a l and other ap p l i c a t i o n s . Nature 211: 481-483. Bakun, A. 1973. Coastal upwelling indices West Coast of North America, 1946-1971. NOAA Tech. Rep. NMFS, SSRF-671 pp. Barber, R.T., and J.H. Ryther. 1969. Organic chelators; factors a f f e c t i n g primary production i n the Cromwell Current upwelling. J. Exp. Mar. B i o l . E c o l . 3_: 191-199. Barber, R.T., R.C. Dugdale, J . J . Maclssac, and R.L. Smith. 1971. Variations i n phytoplankton growth associated with the source and conditioning of upwelling water. Invest. Pesg. 35: 171-193. Barnes, C.A., A.C. Duxbury, and B.A. Morse. 1972. C i r c u l a t i o n and selected properties of the Columbia River e f f l u e n t at sea. Chpt. 3 In: A.T. Pruter and D.L. Alverson (Eds.). The Columbia River Estuary and Adjacent  Ocean Waters. Univ. Wash. Press, Seattle and Washington, 868 pp. Bary, B.M. 1964. Temperature, s a l i n i t y and plankton i n the Eastern North A t l a n t i c and coastal waters of B r i t a i n , 1957. IV. The Species' Relation-ship to the water body; Its r o l e i n d i s t r i b u t i o n and i n s e l e c t i n g and using i n d i c a t o r species. J. Fish. Res. Bd. Can. 21_(1): 183-201. Bary, B.M. 1963a. Temperature, s a l i n i t y and plankton i n the Eastern North A t l a n t i c and coastal waters of B r i t a i n , 1957. I. The ch a r a c t e r i z a t i o n and d i s t r i b u t i o n of surface waters. J. Fish. Res. Bd. Can. 22(3)j^ 789-826. Bary, B.M. 1963b. Temperature, s a l i n i t y and plankton i n the Eastern North A t l a n t i c and coastal waters of B r i t a i n , 1957. I I . The re l a t i o n s h i p s between species and water bodies. J. F i s h . Res. Bd. Can. 20'(4)/I) 1031-1065. 63 Bary, B.M. 1963c. Temperature, s a l i n i t y and plankton i n the Eastern North A t l a n t i c and coastal waters of B r i t a i n , 1957. I I I . The D i s t r i -bution of zooplankton i n r e l a t i o n to water bodies. J. F i s h . Res. Bd. Can. 20(6): 1519-1548. Bary, B.M. 1959. Species of zooplankton as a means of i d e n t i f y i n g d i f f e r -ent surface waters and demonstrating t h e i r movements and mixing. Pac. S c i . 18(1): 14-34. Beattie, J . , C. Bricker, and D. Garvin. 1961. P h o t o l y t i c determination of trace amounts of organic material i n water. Anal. Chem. 30: 1890-1892. Bowen, H.J.M. 1966. Trace elements i n biochemistry. Academic Press, 241 pp. Brinton, E. 1962. The d i s t r i b u t i o n of P a c i f i c Euphausids. B u l l . Scripps. Inst. Oceanogr. 8: 51-270. Brodsky, K.A. 1950. Calanoida of the 'far Eastern seas and the polar basin of the USSR. Zool. Inst. Akad. Nauk., SSSR. Translations, Jerusalem, 440 pp. Campbell, M.H. 1934. The l i f e h i s t o r y and post-embryonic development of the copepods, Calanus tonsus Brady and Euchaeta japonica Marukawa. jJ. B i o l . Ed. Can. 1: 1-65. C a r r i t t , D.E., and J.H. Carpenter. 1966. Comparison and evaluation of cur-r e n t l y employed modifications i n the Winkler method f o r determining d i s -solved oxygen i n seawater; a NASC0 Report. J. Mar. Res. 24: 286-318. Davis, C.C. 1949. The pelagic Copepoda of the Northeastern P a c i f i c Ocean. Univ. Wash. Publ. B i o l . 14: 1-118. Deevy, G.B. 1966. Seasonal v a r i a t i o n s i n the length of copepods i n South P a c i f i c New Zealand waters. Aust. J. Mar. Freshwat. Res. 17: 155-168. 64 Dodimead, A.J., F. Favorite, and T. Hirano. 1963. Review of Oceanography of the Sub-Arctic P a c i f i c Region. International North P a c i f i c F i s h e r i e s Commission B u l l e t i n No. 13. Doe, L.A.E. 1955. Offshore waters of the Canadian P a c i f i c coast. J . F i s h . Res. Bd. Can. 12(1): 1-34. Duxbury, A.C., B.A. Morse, and N. McGary. 1966. The Columbia River E f f l u -ent and i t s d i s t r i b u t i o n at sea. Dept. of Oceanogr. Tech. Rep. No. 156, Univ. Wash. Seattle. Evans, M.S. 1973. The d i s t r i b u t i o n a l ecology of the calanoid copepod Pareuchaeta elongata E s t e r l y . Ph.D. Thesis, I n s t i t u t e of Oceanography and Department of Zoology, Univ. B.C., 105 pp. Fager, E.W., and J.A. McGowan. 1963. Zooplankton species groups i n the North P a c i f i c S c i . 140: 453-460. Gardner, G.A. 1976. The analysis of zooplankton population f l u c t u a t i o n s i n the S t r a i t of Georgia, with emphasis on the re l a t i o n s h i p s between Calanus plumchrus Marakawa and Calanus marshallae Frost. MSc. Thesis, Department of Zoology and I n s t i t u t e of Oceanography, Univ. B.C., 145 pp. Goldberg, E.D. 1957. Biogeochemistry of trace elements. In: J.D. Hedgepeth (ed.) Treatise i n Marine Ecology and Palecology. Vol. I., Ecology. Geol. Soc. Amer. Memoirs 67_: 345-358. Herlinveaux, R.H., and J.P. T u l l y . 1961. Some oceanographic features of Juan de Fuca S t r a i t . J. Fis h . Res. Bd. Can. 18: 1027-1071. Ingelsrud, I., R.J. Robson, and T.G. Thompson. 1936. The d i s t r i b t u t i o n of phosphates i n the seawater of the Northeast P a c i f i c . Univ. Wash. Publ. Oceanogr. 3(1): 1-34. 65 Johnson, R. 1964. Seawater, the natural medium of phytoplankton. 2. Trace metals and chelators, and general discussion. J. Mar. B i o l . Ass. U.K. 44: 87-109. Lewis, A.G., and P.H. W h i t f i e l d . 1974. The b i o l o g i c a l importance of copper i n the sea. A l i t e r a t u r e review. INCRA Contract Report No. 223, 132. pp. Lewis, A.G., P.H. W h i t f i e l d , and A. Ramnarine. 1973. Reduction of copper t o x i c i t y to a marine copepod by sediment extract. Limnol. Oceanogr. 18: 324-327. Lewis, A.G., P.H. W h i t f i e l d , and A. Ramnarine. 1972. Some p a r t i c u l a t e and soluble agents a f f e c t i n g the r e l a t i o n s h i p between metal t o x i c i t y and or-ganism s u r v i v a l i n the Calanoid copepod Euchaeta japonica. Mar. B i o l . 17: 215-221. Lewis, A.G., A. Ramnarine, and M.S. Evans. 1971. Natural chealtors - an i n -d i c a t i o n of a c t i v i t y with the Calanoid copepod Euchaeta japonica. Mar. B i o l . 11: 1-4. Lewis, A.G., and A. Ramnarine. 1969. Some chemical factors a f f e c t i n g the early developmental stages of Euchaeta japonica Marukawa. J. F i s h . Res. Bd. Can. 26: 1347-1362. Lucas, G.E. 1958. External metabolites and p r o d u c t i v i t y . Rapp. Pv. Cons. Inst. Explor. Mer. 144: 155-158. McLaren, I.A. 1965. Some re l a t i o n s h i p s between temperature and egg s i z e , body s i z e , development rate and fecundity of the Copepod Pseudocalanus minutus. Limnol. Oceanogr. 10: 528-538. Menzel, D.W., and R.F. Vaccaro. 1964. The measurement of dissolved organic and p a r t i c u l a t e carbon i n seawater. Limnol. Oceanogr. 9_: 138-142. 66 Morris, B. 1970. Calanoid copepods from midwater trawls i n the North P a c i f i c along 160°W. J. F i s h . Res. Bd. Can. 27: 2297-2321. Patin, S.A. 1973. Some c h a r a c t e r i s t i c s of the occurrence of metals i n the pelagic ecosystem of the ocean. Okeanol. 13: 255-258. Pro v a s o l i , L. 1963. Organic regulation of phytoplankton f e r t i l i t y . In: M.N. H i l l (ed.). The Sea, Vol. I I . Interscience, pp. 165-219, 544 pp. Russel, F.S. 1939. Hydrographical and b i o l o g i c a l conditions i n the North Sea. as indicated by plankton organisms. J. Cons. Int. Explor. Mer. 14(2): 171-192. Russel, F.S. 1936a. A review of some aspects of plankton research. Rapp. et Proc. Verb des Reun. 95: 3-31. Russel, F.S. 1936b. Observations on the d i s t r i b u t i o n of plankton animal i n d i c a t o r s made on Col. E.T. Peel's Yacht "St. George" i n the mouth of the English Channel. J . Mar. B i o l . Ass. U.K. 20: 507-552. Russel, F.S. 1935. On the value of ce r t a i n planktonic animals as indic a t o r s of water movements i n the English Channel and North Sea. J. Mar. B i o l . Ass. U.K. 29(2): 302-332. Sars, G.O. 1925. Copepodes particulierement bathypekigiques provenant des campagnes s c i e n t i f i q u e s . Prince Albert 1-et de Monaco. Res. Camp. S c i . , Fasc. 69. Schubert, J. 1954. Interactions of metals with small molecules i n r e l a t i o n to metal-protein complexes, In: F.R.N. Gurd (Ed.) Chemical s p e c i f i c i t y  i n b i o l o g i c a l i n t e r a c t i o n s . Scott, A. 1909. The copepoda of the Siboga Expedition. Part I. Free-swim-ming, l i t o r a l and semi-parasitic copepoda. Siboga Exped. Monogr. 29a: 1-323. 67 Sewell, R.B.S. 1929. The copepoda i n the Indian Sea. Memoirs of the  Indian Museum 10: 1-407. Slowey, J.F., and D.W. Hood. 1971. Copper, manganese and zinc concentra-tions i n Gulf of Mexico waters. Geochem. Cosmochem. Acta. 35: 121-138. Slowey, J.F., L.M. J e f f r e y , and D.W. Hood. 1967. Evidence for organic Q complexed copper i n seawater. Nature 214: 377-378. Snedecor, G.W. 1956. S t a t i s t i c a l Methods. 5th Ed. Iowa State C o l l . Press, Ames, Iowa. 534 pp. Steeman-Nielsen, E., and S. Wium-Anderson. 1970. Copper ions as poison i n the sea and freshwater. Mar. B i o l . 6_i 93-97. Str i c k l a n d , J.D.H. 1972. Research on the marine planktonic food web at the I n s t i t u t e of Marine resources: A review of the past seven years of work. Oceanogr. Mar. B i o l . Ann. Rev. 10: 349-414. Stri c k l a n d , J.D.H., and T.R. Parsons. 1972. A p r a c t i c a l handbook of seawater analysis. Fish. Res. Bd. Can. B u l l . 167 (Second Edition) 310 pp. Sverdrup, H.U., and R.H. Fleming. 1941. The waters o f f the coast of Southern C a l i f o r n i a , March to July, 1937. B u l l . Scripps. Inst. Oceanogr. 4^: 261-378. Sverdrup, H.U., M.W. Johnson, and R.H. Fleming. 1942. The oceans. Prentice-H a l l , Inc., Englewood C l i f f s , N.J., 1087 pp. Tabata, K., and K. Nishikawa. 1969. Studies on the t o x i c i t y of heavy metals to aquatic organisms and the factors which decrease the t o x i c i t y . V. A t r i a l to decrease the t o x i c i t y of heavy metal ions by the addition of com-plexing agents. B u l l . Tokai Reg. Fish Res. Lab. 58: 225-264. Tanaka, 0., and M. Omaori. 1968. A d d i t i o n a l report on the calanoid copepotio from the Izu region, Part I. Euchaeta and Paraeuchaeta. Publ. Seto Mar. B i o l . Lab. 16: 219-261. 68 T u l l y , J.P. 1942. Surface non-tidal currents i n the approaches to Juan de Fuca S t r a i t . J . F i s h . Res. Bd. Can. .5(4): 389-409. Vervoort, W. 1963. Pelagic Copepoda. Part I. Copepoda of the f a m i l i e s Calanidae up to and i n c l u d i n g Euchaetidae. Atlantidae Rept. 7_: 77-195. Waldichuk, M. 1957. Physical oceanography of the S t r a i t of Georgia, B r i t i s h Columbia. J . Fish. Res. Bd. Can. 14: 321-486. W h i t f i e l d , P.H. 1974. Seasonal changes i n hydrographic and chemical pro-pe r t i e s of Indian Arm and t h e i r e f f e c t on the Calanoid Copepod Euchaeta  japonica. M.Sc. Thesis, U n i v e r s i t y of B r i t i s h Columbia, Department of of Zoology and I n s t i t u t e of Oceanography. 79 pp. W h i t f i e l d , P.H., and A.G. Lewis. 1976. Control of the b i o l o g i c a l a v a i l a -b i l i t y of trace metals to a Calanoid Copepod i n a coastal f j o r d . Est. and Coast Mar. S c i . 4-: 255-266. Williams, P.M. 1968. The ass o c i a t i o n of copper with dissolved organic mat-ter i n seawater. Limnol. oceanogr. 14: 156-158. Williams, P.M. 1968. Stable carbon isotopes i n the dissolved organic mat-ter of the sea. Nature 219: 152-153. Williams, R.F.J. 1953. Metal ions i n b i o l o g i c a l systems. B i o l . Rev. 28: 381-415. Wilson, D.P. 1951. A b i o l o g i c a l difference between natural sea waters. J_. Mar. B i o l . Ass. U.K. 30(1): 1-19. Z i r i n o , A. and S. Yamamoto. 1972. A pH-dependent model for the chemical speciation of copper, zinc, cadmium and lead i n seawater. Limnol. Oceanogr. 17: 661-671. Z i r i n o , A., and M.L. Healy. 1970. Inorganic zinc complexes i n seawater. Limnol. Oceanogr. 15: 956-958. 69 APPENDIX Some disagreement over the name of the bioassay organism i s apparent i n the l i t e r a t u r e . In the S t r a i t of Georgia t h i s species has been studied and r e f e r r e d to by two names: Euchaeta japonica Marukawa, 1921 (Campbell, 1934; Lewis et a l . , 1971, 1972, 1973; W h i t f i e l d and Lewis, 1976) and Pareuchaeta elongata E s t e r l e y , 1913 (Evans, 1973; Gardner, 1976). The genus Euchaeta was established by P h i l l i p p i i n 1882 (Scott, 1909) and the genus Pareuchaeta (Scott, 1909). The d i s t i n g u i s h i n g c h a r a c t e r i s t i c s on which t h i s d i v i s i o n was made were the armature of the second maxilla on the adult female and the structure of the f i f t h l e g on the male. In Euchaeta some of the spines on the ,apeix of the second maxilla are equipped with long spinules; i n Pareuchaeta, the spines are equipped with short s p i n -ules. In the adult male Euchaeta, the t h i r d segment of the exopodite of the f i f t h l e g i s long and spinform; i n Pareuchaeta, i t i s short and rudimentary. Sars (1925, i n Sewell, 1929) added a t h i r d d i s t i n g u i s h i n g feature. In Euchaeta, the accessory setae of the f u r c a l rami are more strongly developed than other f u r c a l setae; i n Pareuchaeta, he found these setae to be quite slender, and form a 'knee j o i n t ' a short distance past t h e i r point of o r i g i n . Vervoort (1963), on examination of t h i r t e e n of the known seventy-six species of Euchaeta and Pareuchaeta, concluded that the separation was not v a l i d . Scott (1909), Sewell (1929), Brodsky (1950) , Tanaka and Omori (1968) examined f i f t y - n i n e of the seventy-six species and concluded that the separ-ation was j u s t i f i e d . As t h e i r conclusion was based on a f a r greater number of representatives the genus Pareuchaeta i t i s here accepted as v a l i d . The structure of the second maxilla (Campbell, 1934), the male f i f t h leg (Campbell, 1934), and the caudal f u r c i (Evans, 1973) i n d i c a t e that the 70 bioassay organism used i n t h i s study belongs to the genus Pareuchaeta. In 1913, Esterley i d e n t i f i e d a new species, Euchaeta elongata, i n the San Diego region. Marukawa, i n 1921, described a new species i n the Sea of Japan which he c a l l e d Euchaeta japonica. Both E s t e r l e y and Marukawa used the name i d e n t i f y i n g c h a r a c t e r i s t i c s . However, the adult female of Marukawa was la r g e r (8 mm i n length vs. 4.13 mm). Brodsky (1959) noted a species i n the Sea of Ohkotsk, the Sea of Japan, the Bering Sea and the northwest P a c i f i c Ocean which he c a l l e d Pareuchaeta japonica. Referring to Esterley's d e s c r i p t i o n he stated t h i s species (japonica) to be i d e n t i c a l to P_. elongata E s t e r l e y . Tanaka and Omori (1968) noted the occurrence of Pareuchaeta elongata i n the Izu region of Japan, and stated that Esterley's elongata and Marukawa' japonica were synonymous. Morris (1970) c o l l e c t e d E_. elongata from the sub-a r c t i c P a c i f i c Ocean, and agreed that elongata and japonica are synonymous species (Evans, 1973, pers. comm.). Some arguments have been made against t h i s on the basis of si z e d i f f e r -ences and q u a l i t a t i v e differences i n the front p a p i l l a and the concavity of the border of the exopod of the f i r s t l e g (Davis, 1949). Evidence e x i s t s that a species may ex h i b i t d i f f e r e n t s i z e s at maturity i n d i f f e r e n t parts of i t s range ( i . e . , Deevy, 1966; McLaren, 1965) and there i s ample evidence of morphological v a r i a t i o n within a species i n d i f f e r e n t parts of i t s range ( i . e . , Brinton, 1962). Based on these arguments (presented by Evans (1973)) and since Pareuchaeta elongata i s the senior name, i t i s used i n the present study. I t should be pointed out, however, that the type material of P_. elongata i s missing (Fleminger, pers. comm. with A.G, Lewis) and that the o r i g i n a l des-c r i p t i o n i s s u f f i c i e n t l y vague to question i d e n t i f i c a t i o n (Lewis, pers. comm. 

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