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The production of planktonic herbivorous food chains in large-scale continuous cultures Brown, Penelope Stevenson 1979

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THE PRODUCTION OF PLANKTONIC HERBIVOROUS FOOD CHAINS IN LARGE-SCALE CONTINUOUS CULTURES by PENELOPE STEVENSON BROWN B . S c , U n i v e r s i t y of V i c t o r i a , 1971  A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY in THE FACULTY OF GRADUATE STUDIES (Department of Zoology, I n s t i t u t e of Oceanography)  He accept t h i s t h e s i s as conforming to t h e , r e q u i r e d standard  THE UNIVERSITY OF BRITISH COLUMBIA August,1979 (c) Penelope Stevenson Brosn,1979  In  presenting  an  advanced  the I  Library  further  for  degree shall  agree  scholarly  by  his  of  this  written  _  thesis  in  at  University  the  make that  it  thesis  purposes  for  partial  freely  permission may  representatives.  ,  financial  gain  Zoology  of  University  of  British  October 10th,  of  of  Columbia,  British  Columbia  1979  for  extensive by  the  is understood  2075 Wesbrook P l a c e V a n c o u v e r , Canada V6T 1W5  Date  for  be g r a n t e d  It  fulfilment  available  permission.  Department The  this  shall  Head  be  requirements  reference copying  that  not  the  of  copying  agree  and  of my  I  this  that  study. thesis  Department or  for  or  publication  allowed without  my  ABSTSACT  Besearch  The examined  scale  large  source of turbulent  the  from  suitability  hastata  of the growth  of  depth  due  and  the  the of  16.8$  per  to a reduced with  with a  environmental  although  an  indicated  that  the  ccnditons. flushing rate  phytoplankton  qigas  primary  culture  communities  variable  flushing rates,  turbulent  upwelling  dynamics of t h e  systems  with  p r i m a r y community  natural  bivalve  was  required  at a  for  conditions. depth  of  temperature at  this  of  0.25/day  provided  growth of  ;oysters,  the two  stocking  the  one  contrast,  monitored from  and  In  experiments,  ranging  two  forcing  concentration  were  were  flushing  growth  levels.  the d y n a m i c s o f at  constant  0.10/day t o  the and  1.00/day, i n  forcing conditions.  were p r e d i c t e d  one-  densities  l i m i t e d the  a t d e n s i t i e s below c o m m e r c i a l  two-stage  systems  t h a t a high  i n t e n s i t y and  of  flushing  ( Chlamys  achieved  for  were  scallops  (0.75/day)  sere  conditions  with  s u r v i v a l of  natural  experimental comparison  of C r a s s o s t r e a  the  light  surface  suitable  of  month  productivity  The  ) and  was deep  p r i m a r y community  results indicated  under  a  u p v e i l i n g systems  the  qigas  chains  using  high  0.7 5/day.  growth and  scallops  food  experiments  dynamics of  The  s t a g e c u l t u r e system  the  to  ( Crassostrea  ).  compared  In  culture  Earsons  cultures  non-turbulent  0.25/day  I.E.  promote  c o n t i n u o u s c u l t u r e system  Maximum r a t e s metre,  seawater t o  f o r the  oysters  hericia  continuous  one-stage  i n terms o f  molluscs,  rate  in  ranging  their  herbivorous  in  investigated  analyzed  planktonic  of  Both  rates  Prof.  production  nutrient-rich rates.  Supervisor:  with  The  reasonable  iii accuracy  using a numerical  determined  p a r a m e t e r s and  growth of the size,  their  source, flushing  oysters density  The  and  results  r a t e of  conditions  was  simulation values also  various  indicated  production  r a t e s a t t a i n e d during the r a t e s measured i n an  of the  examined  1.0/day p r o v i d e d  f o r the  that a the  with  forcing as  primary  a  field  experimentally variables.  function of  communities primary  The  (18%/«eek) location  as  system  most s u i t a b l e  of oysters..  experiment  'optimal'  model  The their  a  food  with  a  environmental  maximum  growth  were g r e a t e r  in B r i t i s h  than  Columbia.  iv TABLE OF CONTESTS  CHAPTER  1. INTRODUCTION  ...................................  1  CHAPTER 2. EXPERIMENTAL F A C I L I T I E S AND METHODOLOGY . . . . . . . . 9 E x p e r i m e n t a l F a c i l i t i e s ................................ 9 One-stage C u l t u r e Experiment H i t h Constant F o r c i n g Conditions.•»..•••...•»•.«••!»••.•»...••••.•••.••««»••• 10 O n e - s t a g e C u l t u r e E x p e r i m e n t s With N a t u r a l F o r c i n g - C o n d i t i o ns ...'«....«...«••..»....«....... .....*..«•...*., 10 Two-stage C u l t u r e E x p e r i m e n t s With N a t u r a l F o r c i n g Conditions ..•.......,............•..•.•.*..»..*.. ...11 13 e t .no do.lo g y ........ m •'•.•....••..•..••••••..•......••••.•*,... • . 12. P h y s i c a l P a r a m e t e r s : L i g h t , T e m p e r a t u r e And S a l i n i t y . . 12 N u t r i e n t P a r a m e t e r s •. ..... ............ ..... ..... ... • .- 1 4 Primary Parameters: Standing Stock, Primary P r o d u c t i v i t y and Community S t r u c t u r e . . . . . . . . . . . . . . . . 15 S e c o n d a r y P r o d u c t i v i t y . . . . . . . . . . . . . . . . . . . . . . . . . . . • - » • , 18 E x p e r i m e n t 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . , 18 Ex p e r i m en t 2 A ......w..*.-*,............... ...:.....*.. . 18 E x per x m en t 2B ........................*........ «w . . . . 19 E xper i BI en t 3ft . . . . . . . . . . . . . . . . . . . . . . . . . . . * . . « « * « « . « . 20 E x p e r i m e n t 3B . . . . . . . . . . . . . . . . . . . . . . . . •..... , .20 E x p e r i m e n t 5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 CHAPTER 3. ONE-STAGE CONTINUOUS CULTURES IK NGN-TURBULENT UPWELLING SYSTEMS WITH CONSTANT FOBC.ING CONDITIONS ...... ,22 D y n a m i c s O f The P r i m a r y C o m m u n i t i e s . . . . . . . . . . . . . . . . . . . . 22 P h y s i c a l E n v i r o n m e n t ...........<•...... ........ . . . . . . . 23 N u t r i e n t C o n d i t i o n s ................................... 23 P h y t o p l a n k t o n Dynamics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 Standing Stock . . . . . . . . . . . . . . . . . . 2 5 P r i m a r y P r o d u c t i v i t y . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 P h y t o p l a n k t o n S t o c k C o m p o s i t i o n . . . . . . . . . . . . . . . . . . ... 27 Growth And S u r v i v a l Of, The S c a l l o p P o p u l a t i o n . . . . . . . . . . 28 CHAPTER 4. ONE-STAGE CONTINUOUS CULTUBES IN NGN-TURBULENT U P » E L X I H G SYSTEMS WITH NATUfiAL FORCING CONDITIONS . . . . . . . . 30 Comparison Of Two Herbivorous Food Chains A t A Low F l u s h i n g R a t e ............ .... • • . . » • . . » • » » . • » . . . . . . . » .. «• »..•. ,-, 31 D y n a m i c s O f The P r i m a r y C o m m u n i t i e s . . . » . . . . . , . . . . , . . - . . ,32 P h y s i c a l E n v i r o n m e n t ... .*».....»•. .. ..."...*.,........... 32 N u t r i e n t C o n d i t i o n s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 E h y t c p l a n k t o n Dynamics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 S t a n d i n g S t o c Jc .»..».>..,. ........................ . 35 .Oxygen ' L e v e l s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 P r i m a r y P r o d u c t i v i t y . . . . . . . . . . ... . 37 C o m p o s i t i o n O f The P h y t o p l a n k t o n Community . . . . . . . . . . . 38 Growth Of The fleriiivores, Crassostrea gigas , During Growth Of The H e r b i v o r e s , C h l a m y s h a s t a t a h e r i c i a , D u r i n g EXP2B . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  40  V  F u r t h e r I n v e s t i g a t i o n Of The O y s t e r Food C h a i n At An Increased-. Her b i v o r e D e n s i t y • . ,., . . . . . . . . . . . . . . , . . . . . 41 D y n a m i c s O f The P r i m a r y Community . . . . . . . . . . . . . . . . . . . . . . 41 P h y s i c a l E n v i r o n m e n t . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ,41 N u t r i e n t Begime . ............................... ....... ,42 P h y t o p l a n k t o n D y n a m i c s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .42 S t a n d i n g S t o c k . . . . . . » . . . . . . . . . . . . . . . . . . . . . . . . . . . .42 Oxygen L e v e l s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43 P r i m a r y P r o d u c t i v i t y ................w............, 44 C o m p o s i t i o n Of The P h y t o p l a n k t o n Community . . . . . . . . . 44 Growth o f t h e O y s t e r s a t a H i g h e r S t o c k i n g D e n s i t y .....,45 ;  F u r t h e r I n v e s t i g a t i o n Of The S c a l l o p Food C h a i n A t An I n c r e a s e d F l u s h i n g B a t e O f The System . . . . . . . . . . . . . . . 46 D y n a m i c s O f The P r i m a r y Community . . . . . . . . . . . . . . . . . . . . . . 46 P h y s i c a l Environment .- ... • . . . . . . . i ,.,.«.>.,, . 46 N u t r i e n t Begime . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ,47 P h y t o p l a n k t o n D y n a m i c s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .47 S t a n d i n g S t oc k ...'..-..«.,.....<....*•. • . . . . . . . . . ....... ,-47 Oxygen L e v e l s . . . . . . . . ...»...»......'....-. ., 48 P r i m a r y P r o d u c t i v i t y . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ,49 C o m p o s i t i o n Of The P h y t o p l a n k t c n Community . . . . . . . . . 4 9 G r o w t h O f The S c a l l o p s A t a H i g h e r F l u s h i n g B a t e .......,49 CHAPTER 5. TBO-STAGE CONTINUOUS COLTUBES*IB TURBULENT UPWELLING SYSTEMS: DYNAMICS OF THE PSIMARY COMMUNITIES AT TWO COMPARATIVE FLUSHING BATES . . . . . . . . . . . . . . . . . . . . . . . . . . . ,51 Dynamics Of The P r i m a r y C o m m u n i t i e s At Two C o m p a r a t i v e F l u s h i n g B a t e s .....», .......«.'......-.... ,52 P h y s i c a l E n v i r o n m e n t . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52 N u t r i e n t Regime .......................................53 P h y t o p l a n k t o n D y n a m i c s .... ....... . . . . . . . . . . . . . . . . . . . . .54 S t a n d x n g Sitock . • • . . • • • • . • • • . • • . . . . . . . . . . . . . . . . . . . » . 54 Pigment R a t i o s . . . . . . . . . . . . . . . . .... 56 Oxygen L e v e l s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56 P r i m a r y P r o d u c t i v i t y . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58 E s t i m a t i o n Of Parameters F o r A Primary P r o d u c t i v i t y Model .... . . . . . . . . . . . . . . . . . . . . . . . . . . . .61 C o m p o s i t i o n O f The P h y t c p l a r k t o n Community . . . . . . . . . 63  CHAPTER 6. 3HO-STAGE CONTINUOUS COLTUBES OF I1ANKTONIC HEBBIVOJOUS FOOD CHAINS: DYNAMICS OF TJTE PEIMABY COMMUNITIES AT VARIABLE FLUSHING BATES ...................................65 Dynamics O f The P r i m a r y Communities At V a r i a b l e F l u s h i n g B a t e s D u r i n g The E x p e r i m e n t a l P e r i o d . . . . . . . . . . . . . . . . 66 P h y s i c a l E n v i r o n m e n t . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67 N u t r i e n t Begime . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68 P h y t o p l a n k t o n D y n a m i c s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70 S t a n d i n g S t o c k . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70 P i g m e n t B a t i o s ...................................... 71 Oxygen L e v e l s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72 P r i m a r y P r o d u c t i v i t y . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73 E s t i m a t i o n Of P a r a m e t e r s F o r A P r i m a r y P r o d u c t i v i t y Model . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ,76 C o m p o s i t i o n O f The P h y t o p l a n k t o n Community . . . . . . . . . 77  vi  CHAPTER 7. TWO—ST AGE CONTINUOUS CULTURES OF PLANKTGN.IC HERBIVOROUS FOOD CHAINS: GROWTH OF THE HEBE.IVORES . . . . . . . . 80 E x p e r i m e n t a l D e s i g n . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ,80 E x p e r i m e n t a l R e s u l t s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81 E n v i r o n m e n t a l C o n d i t i o n s D u r i n g The O y s t e r Growth ; Experiments ............................»»......*....,81 O y s t e r Growth As A F u n c t i o n Of The E x p e r i m e n t a l F a c t o r s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85 CHAPTER 8. ANALYSIS OF THE RESULTS USING A SIMULATION MO DE L . ., .v. . . . ......... .......................... . 89 Estimation Of T h e P h y s i o l o g i c a l P a r a m e t e r s D u r i n g The Continuous Cultures . . . . . . . . . . . . . . . . . . . . . . . . . . . 89 S i m u l a t i o n Of The P h y t o p l a n k t o n Dynamics .,....,.,.,.-,.92 D e s c r i p t i o n O f The Model . . . . . . . . . . . . . . . . . , 9 2 Simulation Results . . . . . . . . . . . . . . . . . . . . w . . , 9 5 CHAPTER  9.  COMPARATIVE RESULTS  CHAPTER  10. SUMMARY  DISCUSSION OF THE EXPERIMENTAL ......99  AND CONCLUSIONS  107  T ABLES  «. ......... ...,110  FIGURES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ,149  BIBLIOGRAPHY .  APPENDICES .  ,. , i , . . . . v . . . . .  .. .  ., , .  ,  253  . . . . . . . . . . v . ' . . , . , - , . . . - . . . ,260  vxx  LIST OF TABLES  Table Table Table Table Table Table Table Table Table Table Table Table Table Table Table Table Table Table Table Table Table Taijle Table Table Table Table  1. B e l e v a n t c u l t u r e s t u d i e s o f m a r i n e p r o d u c t i o n 2. D e s c r i p t i v e s t a t i s t i c s f o r E x p e r i m e n t 2A 3. D e s c r i p t i v e s t a t i s t i c s f o r E x p e r i m e n t . 2B 4. O y s t e r g r o w t h d u r i n g E x p e r i m e n t 2A 5. S c a l l o p growth d u r i n g E x p e r i m e n t 2.B 6. D e s c r i p t i v e s t a t i s t i c s f o r E x p e r i m e n t 3A 7. O y s t e r g r o w t h d u r i n g E x p e r i m e n t 3A 8. D e s c r i p t i v e s t a t i s t i c s f o r E x p e r i m e n t 3B 9., S c a l l o p g r o w t h d u r i n g E x p e r i m e n t 3B 10. C o r r e l a t i o n between f o r c i n g v a r i a b l e s d u r i n g EXP4 11.. D e s c r i p t i v e s t a t i s t i c s f o r E x p e r i m e n t 4A 12. D e s c r i p t i v e s t a t i s t i c s f o r E x p e r i m e n t 4B 13. A n a l y s i s o f v a r i a n c e d u r i n g E x p e r i m e n t 4A 14. A n a l y s i s o f v a r i a n c e d u r i n g E x p e r i m e n t 4B 15. P r o d u c t i v i t y component a n a l y s i s f o r E x p e r i m e n t s 16. D e s i g n f o r t h e t w o - s t a g e c u l t u r e e x p e r i m e n t s 17. N u t r i e n t e n r i c h m e n t e x p e r i m e n t r e s u l t s 18. D e s c r i p t i v e s t a t i s t i c s f o r E x p e r i m e n t 5A 19. D e s c r i p t i v e s t a t i s t i c s f o r E x p e r i m e n t 5B 2 0 . A n a l y s i s o f v a r i a n c e d u r i n g E x p e r i m e n t 5A 2 1 . A n a l y s i s o f v a r i a n c e d u r i n g E x p e r i m e n t 5B 22. P r o d u c t i v i t y r e s u l t s d u r i n g E x p e r i m e n t 5 2 3 . P r o d u c t i v i t y component a n a l y s i s f o r E x p e r i m e n t 5 24. E n v i r o n m e n t a l c o n d i t i o n s i n t h e h e r x i v o r e t a n k s 25. Growth o f o y s t e r s i n t h e t w o - s t a g e c u l t u r e 2 6 . Growth o f o y s t e r s a s a f u n c t i o n o f FJJP, S I Z E and DENS i n t h e t w o - s t a g e c u l t u r e T a b l e 27. N o n - l i n e a r LSF o f t h e *P v e r s u s I * d a t a f o r t h e TANH and SMITH models  p.110 p..111. p. 112 P-113 p. 114 p.115 p. 116 p.117 p.118 p.119 p. 120 P«122 p. 124 p.126 p. 128 p.129 p.130 p. 131 p.132 p.133 p.135 p. 137 p.138 p.139 p.140 p. 143 p.144  viii  LIST OF  Figure Figure Figure Figure Figure Figure Figure Figure Figure Figure Figure Figure  1. , 2.< 3. 4., 5. 6, 7. 8. 9., .10. ., 11., 12.  F i g u r e 13. . F i g u r e 14., F i g u r e 15., F i g u r e 16. F i g u r e 17. F i g u r e 18. F i g u r e 19. F i g u r e 20. F i g u r e 21. F i g u r e 22. F i g u r e 23. F i g u r e 24. F i g u r e 25. Figure Figure Figure Figure Figure Figure Figure  26. 27. 28. 29. 30. 31. 32.  F i g u r e 33. F i g u r e 34. , F i g u r e 35., F i g u r e 36. F i g u r e 37. F i g u r e 38. , F i g u r e 39.  E x p e r i m e n t a l f a c i l i t i e s f o r E x p e r i m e n t 1. p.149 D u p l i c a t e t a n k s y s t e m s f o r E x p e r i m e n t s 2 and 3, p.150 E x p e r i m e n t a l f a c i l i t i e s f o r E x p e r i m e n t s 2 and 3. p.151 Experimental f a c i l i t i e s for Experiments., p. 152 T w o - s t a g e c u l t u r e s y s t e m s f o r E x p e r i m e n t 5. p.153 N i t r a t e c o n c e n t r a t i o n d u r i n g E x p e r i m e n t 1A. P-154 N i t r a t e c o n c e n t r a t i o n d u r i n g E x p e r i m e n t LB. p.155 P h y t o p l a n k t o n s t o c k d u r i n g E x p e r i m e n t 1A. p.156 P h y t o p l a n k t o n s t o c k d u r i n g E x p e r i m e n t IB. P-157 P r i m a r y p r o d u c t i v i t y d u r i n g E x p e r i m e n t 1A., p.158 P r i m a r y p r o d u c t i v i t y d u r i n g E x p e r i m e n t LB. p.159 Primary p r o d u c t i v i t y (standardized ) during E x p e r i m e n t 1B p. 160 S o l a r r a d i a t i o n d u r i n g E x p e r i m e n t 2., p.161 T e m p e r a t u r e d u r i n g E x p e r i m e n t 2A. p.162 Temperature d u r i n g Experiment 2E. , p.163 N i t r a t e c o n c e n t r a t i o n d u r i n g E x p e r i m e n t 2A. P-164 N i t r a t e c o n c e n t r a t i o n d u r i n g E x p e r i m e n t 2B. P-165 P h y t o p l a n k t o n s t o c k d u r i n g E x p e r i m e n t 2A. , p.166 P h y t o p l a n k t o n s t o c k d u r i n g E x p e r i m e n t 2B. p.167 Oxygen c o n c e n t r a t i o n d u r i n g E x p e r i m e n t 2A. p.168 Oxygen c o n c e n t r a t i o n d u r i n g E x p e r i m e n t 2B. p.169 P r i m a r y p r o d u c t i v i t y d u r i n g E x p e r i m e n t 2A., p.170 P r i m a r y p r o d u c t i v i t y d u r i n g E x p e r i m e n t 2B. p.171 Primary p r o d u c t i v i t y (standardized) during E x p e r i m e n t 2A. p. 172 Primary p r o d u c t i v i t y (standardized) d u r i n g E x p e r i m e n t 2B. p.173 S o l a r r a d i a t i o n d u r i n g E x p e r i m e n t 3A. p.174 T e m p e r a t u r e d u r i n g E x p e r i m e n t 3A. , p.175 N i t r a t e c o n c e n t r a t i o n d u r i n g E x p e r i m e n t 3A. , p. 176 P h y t o p l a n k t o n s t o c k d u r i n g E x p e r i m e n t 3A. P-177 Oxygen c o n c e n t r a t i o n d u r i n g E x p e r i m e n t 3A. p.178 P r i m a r y p r o d u c t i v i t y d u r i n g E x p e r i m e n t 3A. p.179 Primary p r o d u c t i v i t y (standardized) during E x p e r i m e n t 3A. , p. 180 S o l a r r a d i a t i o n d u r i n g E x p e r i m e n t 3B. p. 181 Temperature d u r i n g Experiment 3£. p.182 N i t r a t e c o n c e n t r a t i o n d u r i n g E x p e r i m e n t 3B., P-183 P h y t o p l a n k t o n s t o c k d u r i n g E x p e r i m e n t 3B. p.184 Oxygen c o n c e n t r a t i o n d u r i n g E x p e r i m e n t 3B., p.185 P r i m a r y p r o d u c t i v i t y d u r i n g E x p e r i m e n t 3B. p.186 Primary p r o d u c t i v i t y (standardized) during E x p e r i m e n t 3B. p. 187 F o r c i n g c o n d i t i o n s d u r i n g Experiment 4 p.188 T e m p e r a t u r e d u r i n g E x p e r i m e n t 4., p. 189 N i t r a t e c o n c e n t r a t i o n d u r i n g E x p e r i m e n t 4. p.190 P h y t o p l a n k t o n s t o c k d u r i n g E x p e r i m e n t 4.. P-191 CHLB:CHLA r a t i o d u r i n g E x p e r i m e n t 4. p.192 C a r o t e n o i d : C H L A r a t i o d u r i n g E x p e r i m e n t 4., p.193 Oxygen c o n c e n t r a t i o n d u r i n g E x p e r i m e n t 4. p.194 G r o s s p r i m a r y p r o d u c t i v i t y d u r i n g E x p e r i m e n t 4. p.195 E e s p i r a t i o n r a t e d u r i n g E x p e r i m e n t 4. p.196 Net p r i m a r y p r o d u c t i v i t y d u r i n g E x p e r i m e n t 4. p.197 ;  Figure Figure Figure Figure Figure Figure Figure Figure Figure Figure  40.. 41. 42. 4 3. 44., 45. 46. , 47. 48. 49.  FIGUBES  ix  F i g u r e 50.  Net p r i m a r y p r o d u c t i v i t y ( d a i l y ) d u r i n g E x p e r i m e n t 4. F i g u r e 51. Gross primary p r o d u c t i v i t y (daily) during Experiment,4. F i g u r e 52. G r o s s p r i m a r y p r o d u c t i v i t y (standardized) d u r i n g E x p e r i m e n t 4. F i g u r e 53. Net primary p r o d u c t i v i t y (standardized) d u r i n g E x p e r i m e n t 4. F i g u r e 5 4. Respiration rate (standardized) diiring E x p e r i m e n t s . F i g u r e 5 5 . , E s t i m a t e s o f ALPHAC d u r i n g E x p e r i m e n t 4 F i g u r e 56. „ E s t i m a t e s o f ALPHAG d u r i n g E x p e r i m e n t 4. F i g u r e 57. C o u l t e r c o u n t s on Day 6 o f E x p e r i m e n t 4A. , F i g u r e 5 8 . , C o u l t e r c o u n t s on Day 9 o f E x p e r i m e n t 4A. , F i g u r e 5 9 . C o u l t e r c o u n t s on Day 15 o f E x p e r i m e n t 4A. F i g u r e 6 0 . , C o u l t e r c o u n t s on Day 21 c f E x p e r i m e n t 4A. F i g u r e 6 1 . , C o u l t e r c o u n t s on Day 27 c f E x p e r i m e n t 4 A. F i g u r e 62., F o r c i n g c o n d i t i o n s d u r i n g E x p e r i m e n t 5. F i g u r e 6 3 . T e m p e r a t u r e d u r i n g E x p e r i m e n t 5. F i g u r e 64., N i t r a t e c o n c e n t r a t i o n d u r i n g E x p e r i m e n t 5. F i g u r e 6 5 . P h y t o p l a n k t o n s t o c k d u r i n g E x p e r i m e n t 5. F i g u r e 6 6 . C H L £ A : C B L A r a t i o d u r i n g E x p e r i m e n t 5., F i g u r e 67. C a r o t e n o i d i C H L A r a t i o d u r i n g E x p e r i m e n t 5. F i g u r e 6 8 . Oxygen c o n c e n t r a t i o n d u r i n g E x p e r i m e n t 5., F i g u r e 69. G r o s s p r i m a r y p r o d u c t i v i t y d u r i n g Experiment 5 F i g u r e 70. Gross primary p r o d u c t i v i t y ( d a i l y ) during Experiment 5 F i g u r e 71. B e s p i r a t i o n r a t e d u r i n g Experiment 5 F i g u r e 7 2 . Net p r i m a r y p r o d u c t i v i t y d u r i n g Experiment 5 F i g u r e 7 3. Net p r i m a r y p r o d u c t i v i t y ( d a i l y ) d u r i n g Experiment 5 F i g u r e 74. Gross primary p r o d u c t i v i t y (standardized) during Experiment 5 F i g u r e 75. B e s p i r a t i o n r a t e (standardized) d u r i n g Experiment 5 F i g u r e 76. N e t p r i m a r y p r o d u c t i v i t y (standardized) during Experiment 5 F i g u r e 7 7 . E s t i m a t e s o f ALPHAC d u r i n g E x p e r i m e n t 5 F i g u r e 7 8 . E s t i m a t e s o f ALPHAG d u r i n g E x p e r i m e n t 5 F i g u r e 7 9 . C o u l t e r c o u n t s on Day 6 o f E x p e r i m e n t 5A. F i g u r e 8 0 . C o u l t e r c o u n t s on Day 9 o f E x p e r i m e n t 5A. F i g u r e 8 1 . C o u l t e r c o u n t s on Day 21 o f E x p e r i m e n t 5A. F i g u r e 82. C o u l t e r c o u n t s on Day 24 o f E x p e r i m e n t 5A. F i g u r e 8 3 . , C o u l t e r c o u n t s on Day 36 o f E x p e r i m e n t 5A. F i g u r e 8 4 . C o u l t e r c o u n t s on Day 39 c f E x p e r i m e n t 5A. F i g u r e 85.. C o u l t e r c o u n t s on Day 6 o f E x p e r i m e n t 5B. F i g u r e 8 6 . C o u l t e r c o u n t s on Day 9 o f E x p e r i m e n t 5B. F i g u r e 8 7 . C o u l t e r c o u n t s on Day 21 o f E x p e r i m e n t 5B. F i g u r e 88. C o u l t e r c o u n t s on Day 24 c f E x p e r i m e n t 5B. F i g u r e 8 9 . C o u l t e r c o u n t s on Day 36 c f E x p e r i m e n t 5B. Figure,90. C o u l t e r c o u n t s on Day 39 o f E x p e r i m e n t 5B. F i g u r e 91. Temperature i n two-stage o y s t e r c u l t u r e s F i g u r e 92. C h l o r o p h y l l a i n two-stage o y s t e r c u l t u r e s F i g u r e 9 3 . Oxygen l e v e l s i n t w o - s t a g e o y s t e r c u l t u r e s  p.198 p.199 p.200 p. 201 p. 202 p. 203 p.204 p. 205 p.206 p.207 p.208 p. 209 p.210 p.211 p.212 p.213 P»214 p.215 P-216 P-217 p.218 P-219 p.220 P»221 p.222 P* 223 p.224 p.225 p.226 p.227 p. 228 p.229 p.230 p.231 p. 232 p.233 p. 234 p.235 p.236 p. 237 p.238 p.239 p.240 p.241  X  F i g u r e 94. F i g u r e 9 5. Figure  96.  Figure  97,  Figure  98, .  F i g u r e 99. Figure  100.  Figure  101.  Figure  102.  F i g u r e 103;  Plate I., Plate I I .  Growth o f o y s t e r s a s a f u n c t i o n of DENS i n the t w o - s t a g e c u l t u r e Growth o f o y s t e r s as a f u n c t i o n o f S I Z E i n the two-stage c u l t u r e 'P v e r s u s ' I * c u r v e s a s a f u n c t i o n o f TEMP and N03 Simulated phytoplankton stock during E x p e r i m e n t 5B: Bun 1 Simulated phytoplankton stock during E x p e r i m e n t 5B: Run 2 Simulated phytoplankton stock during E x p e r i m e n t 5B: Bun 3 Simulated phytoplankton stock during E x p e r i m e n t 5E: Sun 4 Simulated phytoplankton stock during E x p e r i m e n t 5A: Bun 1 Simulated phytoplankton stock during E x p e r i m e n t 5&: Bun 2 Simulated phytoplankton stock during E x p e r i m e n t 5B: Bun 1 w i t h GRAZE H e r b i v o r e t a n k s i n two-stage experiments A r t i f i c i a l c u l t c h i n two-stage experiments  p.242 P-243 p,244 P-245 p.246 p.247 p.248 p. 249 p. 250 p.251 p.252 p.252  xi  ACKNOWLEDGEMENT I would l i k e t o a c k n o w l e d g e , w i t h t h a n k s , t h e s u j p c r t and a s s i s t a n c e o f irany p e o p l e , who c o n t r i b u t e d t o t h e c o o p l e t i c c o f t h i s s t u d y . I would a l s o l i k e t o a c k n o w l e d g e t h e N a t i o n a l R e s e a r c h C o u n c i l c f Canada f o r p r o v i d i n g f i u a r c i a l s u p j e r t with Post-Graduate S c h o l a r s h i p s . I would f i r s t l i k e t o t h a n k my s u p e r v i s o r , Dr. I . E . Pars e n s , f o r h i s i n s p i r a t i o n i n the i n i t i a t i o n of t h i s research, c r i t i c i s m o f i t s d e s i g n and f i n a n c i a l s u p p o r t f o r t h e e x p e r i m e n t a l operations, i n c l u d i n g the invaluable assistance c f B i l l L i during the c n e - s t a g e c u l t u r e e x p e r i m e n t s , and A n g e l a N o r t o n and C a r c l e Eawden o f t h e I n s t i t u t e c f O c e a n o g r a p h y , U.B.C., f o r a n a l y s i s o f n u t r i e n t s a m p l e s . Many o t h e r p e o p l e a t t h e I n s t i t u t e were a l s o most h e l p f u l , I would a l s o l i k e t o a c k n o w l e d g e the c o - o p e r a t i o n o f t h e F i s h e r i e s and M a r i n e S e r v i c e , Department o f t h e E i v i r c n a e n t , for providing experimental f a c i l i t i e s a t the P a c i f i c Envirornsent I n s t i t u t e , West V a n c o u v e r , and i n p a r t i c u l a r , t h a c k t h e many s c i e n t i s t s and t e c h n i c i a n s vhc p r o v i d e d v a l u e d s c i e n t i f i c a r d m o r a l s u p p o r t w h i l e t h e e x p e r i m e n t s were b e i n g c o n d u c t e d . I am a l s c most g r a t e f u l t o Dr. J . i . C . T o n s l i n s c n , F a c u l t y o f Commerce and B u s i n e s s A d m i n i s t r a t i o n , U . B . C , f o r h e l p f u l d i s c u s s i o n s c n s y s t e m s r e s e a r c h , and t o C S . C , s i H i e f o r h e r e x p e r t a d v i c e i n the d e s i g n o f a c o m p u t e r d a t a r a s e and i n the a n a l y s i s o f t h e d a t a . F i n a l l y , I would l i k e t o thank my c o m m i t t e e f o r h e l p f u l d i s c i s s i o n s d u r i n g t h i s s t u d y : Dr. I . E . P a r s e r s , D r . A.G. L e s i s , and Dr. F.J..B. T a y l o r o f t h e I n s t i t u t e o f O c e a n o g r a p h y ; Dr. T.H. C a r e f o o t o f t h e Department o f Z o o l o g y ; and Dr. D.K. Farmer o f t h e I n s t i t u t e c f Ocean S c i e n c e s , P a t r i c i a Eay, E r i t i s h C o l u m b i a .  1  CHAPTER  Maximizing organic  1.,  INTRODUCTION  production  e s s e n t i a l component o f e f f e c t i v e and  D i c k i e , 1970; S a i l a ,  1970),  and  Bierman e t a l . ecosystems,  ,  as  e s t u a r i n e environments Byther,  1969),  indicate  primary,  secondary,  of  ecosystems  these  multidimensional the  production  the  models  and  to  predictable, model; a r e  with  natural  or  particularly estimated  inherent artificial  borrowed  time:  space  and  and  studies  explanatory  varying  complexity  food  chain or  Wiegert  et a l .  However a t t h i s  ncn-linearities,  are  values f o r the  frcm  complex  ecosystem  perturbations  i f parameter  or  1961;  (1971),  (1973) r e s p e c t i v e l y .  1971) and  Schleske,  particular  Dugdale  productive  Extensive  experiments  have d e s c r i b e d  , 1975;  and o r g a n i z a t i o n o f  communities.  field  o f i n t e r a c t i o n s , with  responses  the production  and o r g a n i z a t i o n o f t h a t  (1973) a n d C a p e r on  of  (Pillay,  et a l .  highly  and  an  (Paloheimo  (Cushing,  importance  and t e r t i a r y using  Toro of  Odum  i s  development  upwelling  1962;  both  web, e x e m p l i f i e d by i a l s h  level  of  (Teal,  v a r i a b l e s i n determining  (Di  Observations  areas  management  aguaculture  control  1974).  such  fisheries  1971),  eutrophicaticn  i n a q u a t i c ecosystems  less  original  the l i t e r a t u r e  (Saila,  1973) Two  complementary  situation of  are,f i r s t ,  basis,  ecosystems,  with  approaches  t o reduce the complexity  t h e n a t u r a l ecosystem  trophic  as  research  and  by  secondly,  known f o r c i n g  c o n t r o l parameters.,  initially to  improve  investigate  case  this  and c o n n e c t i v i t y  »subsystemizing*  c o n d i t i o n s using  In t h e f i r s t  to  time  on  a  controlled and  space  f o r example, a n a l y s i s  2  of  phytoplaivk t o n i c  subsystems  eatrophic  environments  Van  1968;  Dyne,  organization 1972),  rate 1971),  McLaughlin,  in  the  or to  c a s e , •;• c o n t r o l l e d  1975;  Subba Hao,  1970;  system  in  terms  time,  organic  space, as  and  1973)  and  expressed 1969; In  as  the  Cassin  and  the  production  trophic levels,  o f t i m e and  (Patten  McAllister st a l . , ,  ( P i a t t , 1972)••  experiments of  naturally  Smayda,  (Dickman,  space  o p e r a t i o n a l s c a l e , number of  control  to  of  production  ,  ;  specifically  of  studies  emphasize  et a i ,  and  seme r e l a t i n g  flushing  which  Winter  (Piatt  include  second  have  and  varied  operational  outlined later  in  this  chapter. Conditions have  been p r o p o s e d  (1962),  and  and  the  of  controlled and  considered  by  al.  Mullin The  Bella  in  hypothesis enhanced  by  provided  examine  was  that  using  the  as  turbulent  addition to  £t  the  the  mixing  these  field  approaches  systems  have  (1973a),  a l .,  rate  production  basis f o r the in  on  to been  Grenney  (1977)  ( 1 9 7 4 ) , and  for Steele  levels.  flushing  production  production  such  level Nielsen  focusing  (1974), S t e e l e  trophic  the  primary  theoretical  Piatt  determining  marine ecosystems, to  of  by  Grenney e t a l .  (1973),  Phillips  the  (1962), Steeman  planktcnic  (1970),  Boss by  more of  at  (1972)  considerations  organization  and  Menzel  water c o l u m n , . In  importance  strategy  and  Parsons  (1977) f o r h i g h e r  considerations  production  experiments,  (1973b),  phytoplankton and  Steele  spatial  depth of the  production  et  by  T a k a h a s h i and  importance and  f o r maximizing  and and  spatial  structure  present  planktonic  i n b i v a l v e feed  of  research  systems..  chains  of  could  a deep n u t r i e n t - r i c h s o u r c e o f s e a w a t e r and  The be an  optimal  flashing rate  and  s p a t i a l structure  maximize  productivity.  duplicate  c o n t r o l l e d e c o s y s t e m s were d e s i g n e d  operational 1.  a  In  order  continuous  flushing  rate  seairater  either  could  culture  be'  upwelled  mixed.  2.  nutrient-rich  deep  3.  a  of  relatively  a choice light  and  5. / u t i l i z a t i o n by  two-stage  with  of  the  placing  the  culture  this  water  seawater  the  column primary  production herbivores  system,  the  and  to  the or  enhance  herbivore  controlled  production  and  to  forcing  conditions  system r e s p o n s e ,  in situ  to  or,  simulate  system,  and  , or  primary  the The  various  rates  a  one-stage  expansion to production  objectives first  organization  mathematical  flushing  as  a  tank  tanks.  of  of  the  of  objective  flushing rates  analyze numerically  deterministic The  1 metre) t o  community.  systeii  systems f o r a s i g n i f i c a n t period  techniques..  (about  conditions  with the  were e s s e n t i a l l y t w o - f o l d .  month) , a  of  environmental  p h y t o p l a n k t o n i c community a t  one  source  i n which  field.  i n t o a s e t of  Using  upwelling  following  minimum o f t u r b u l e n c e  temperature; t o s i m p l i f y the  i n the  examine  with the  controlled,  between c o n t r o l l e d , c o n s t a n t  production  research  a  l i m i t a t i o n o f .the  fluctuating  feeding  hypothesis,  system  and  with  shallow  naturally  culture  this  to  productivity.  minimize l i g h t 4.  tank  varied  artificially  primary  test  ecosystem  capabilities:  large  a  to  c f the  time  using  ranged from  0.10  was  to  a  natural  in  turbulent  (greater  ecosystem  model  the  than  responses simulation  day-1  to  1.0  4  day-1 is  during  the e x p e r i m e n t s s i n c e the  enhanced  represent The the  at  Populations  heric la  )  specific  substations and  and  effect  both  of o y s t e r s , and  1  depths on  harvestable food  the  using  the  per  commercial  of  be  cultch),  production.  determined a s  to  The  system.  The  phytoplankton, were a n a l y z e d Examples  of  the  hastata  was  commercially level  heterogeneity In the  the of  of  gigas  the  (with  cultch  were eight  used  in  ycung o y s t e r s c o u l d  the f o l l o w i n g f a c t o r s :  urea  in  two-stage  flushing  i n the  size  r a t e of  temperature, standing  o x y g e n , ammonia and  tanks.  monitored  efficient  cultch  o y s t e r s , and  the  production  Crasscstrea  simulate  a f u n c t i o n of  dynamics  ), in  Chlamys  produce a  controlled.  production  the o y s t e r s , d e n s i t y of the  two-stage  B i v a l v e s were c h o s e n  constructed  designed  and  community  spatial  juvenile  artificially  growth o f  primary  growth o f  examined  impoundments.,  i n the  culture  experiment,  they  c a g e s , were p o s i t i o n e d a t  experimentally  then  (  ecologically  c h a i n . , Furthermore,  and  i n net  primary  a t a low,  1972)  ( Crassostrea gigas  herbivores s i n c e they  resource  phytoplankton  resulting  one-stage  scallops  h e r b i v o r e s c o u l d be  oysters  mariculture  the one-stage c u l t u r e experiments..  the experimental  the  in  of  Parsons,  t o examine the  which were c o n f i n e d  growth  during  for small  populations  of c o m m e r c i a l c u l t c h  Their  (Brown and  s e c o n d o b j e c t i v e was  cultures.  as  rates  attainable flows  herbivore  form  these  growth  stock  herbivore  the of  tanks,  as c o v a r i a t e s . , of  studies  * A large oyster h a l f ^ s h e l l s t r i n g i n g frcm r a f t s . „  pertinent  with  to  this  research,  oyster spat attached  which  to i t , f o r  5  b a s i c a l l y includes productive categorized  i n Table  First,  there  example,  Lummi I n d i a n on  the  and  salaouids. seawater  Tribal  A  at  25  °C .  day  Hood, 1970)  bacterial  nine  Of  from  environment,  the  Halone e t a l .  with The  and  ,  Malone study duration  The  populations,  favoured  the  species.  ft  be  and  reducing  Parsons,  maximization  raising  nutrient-rich beneficial  high  surface  notable  in  their  operational coatrol  in  time  c u l t u r e s t u d i e s , two  and  in  nutrient-rich a semi-tropical  temperatures greater value  a sub-optimal which  with  than  only  herbivore  a  controlled  rather to  examined  than  the  due  on  the  Crassostrea research  upwelling  s e a w a t e r and  of primary  ecosystem  present  simulated  (  diet  concentrated  that their  edulis  stability  (Baab  a phytoplariktcn population  experiment  1972)  utilized  r a t e s of n u t r i e n t - r i c h and  for rearing  more  are  conducted  determined  growth o f O s t r e a  flushing  even  the  relies  for  deep  could  Baab s t u d y ,  preliminary  supply diets  i s of l i m i t e d  to i t s morphology.  the  food  1975)  were  i s probably  on  m a r i c u l t u r e system  average experimental  mollusc  level.,  impoundment o f  utilizing  four continuous  depth  field  have  levels.  C h a e t o c e r o s ) which  various  tidal  large s c a l e experiments  experimental  a»d  large-scale  as t h e  impoundment  productivity,  , 1973;  seawater  (Brown  a  o f t r o p h i c a n a l y s i s and  space. al.  marine ecosystems which  E n t e r p r i s e . ... T h i s  ( S h i e l s and  variability  four  few  controlled  marine  Secondly,  ca.  keen  are  observations.  a supplement t o the c o m m e r c i a l  t e m p e r a t u r e s and  et  following  however* i s t h e 7 50 a c r e  enhancing  and  the  natural surface plankton  molluscs  by  with  have  been e x p e r i m e n t a l l y One  I  marine p l a n k t e n i c ecosystems,  the  communities.  at  effect  6  Thirdly, has  been  an o r g a n i c  used  dilated  sewage,  i n a s e r i e s of medium-scale c o n t i n u o u s  cultures  investigated  by  specialized  system  source of  Dunstan  and  nutrients,  Tenore,  and  f o r eutrophication  provides  a  c o n t r o l and  useful  mariculture  production. Fourthly, included  as  insights  into  limited  examples  been  Eppley  either  et a l . , Harrison  Finally,  mostly  studies  »  ,  although  of  in  a  and  optimization,  ,  Table laboratory  I  are  provide they are  studies  represents  i s  and Myer, Davis,  (  elucidate and  light  and D y e r , 1965; Caperon,  1967;  1972; D a v i s e t a l . 1976;  studies i n Table (ialker  Conway, I have  1977). utilized  and Z a h r a d n i k ,  although  has t h e d i s a d v a n t a g e  in  1976;  terms  of  of providing  only  production..  on t h e the  (Eppley  in  techniques  to  temperature  chemostat  1974),  i s not intended  molluscs, I  a  control  Barber,  or two-dimensional  bivalve  in  1976;  t h i s system  small-scale  nutrient,  , 1 9 7 1 ; Caperon  f o r operational  Kirby-Smith  a  turbidostat  1964) o r  et a l . ,  continuous c u l t u r e  on  two o f t h e s m a l l - s c a l e  raceway  Table  production  research,  principles  Maddux a n d J o n e s ,  areal  continuous culture  c o n t r o l l e d e x p e r i m e n t s which  maximizing  applied  limitation,  a  of  I n plankton  physiological  1973;  small-scale  i n s c a l e f o r a p p l i c a t i o n s t o mass c u l t u r e p r o d u c t i o n  hatcheries. have  five  a s a complete  growth  herbivores a  sample  of  marine  used of  » f o r a discussion o f the continuous T a y l o r (1960) and Oppenheimer ( 1 S 6 6 ) .  listing  of f i e l d o r  phytoplankton. or  i n this research., studies  culture  which.  technique,  What have  see  7  e x p e r i m e n t a l l y acknowledged of  operation,  parameters, the  trophic  herbivores.,  and  Other  have  ( L o o s a n o f f and using  rafts  ranged  D a v i s , 1963;  Imai,  in  the  and  and  on p h y s i o l o g i c a l f a c t o r s  (1970)  which  which  as  research  presented  provide  various  continuous culture were  in  of seed  bivalves  ( P a r s o n s , 1974)  affect  to  Some  have  the growth  Dunstan  of  Baggaley (1973b,c)  o f the t y p e o f  relationship  this  comparisons  flushing  experiments, conducted  c o n d i t i o n s of l i g h t ,  between  between  the  continuous  and  of  this  1  food  and  two-stage  system.  Five  three to ten  weeks  The  were  forcing  essentially  2 t o 5 were  conducted  i n o r d e r t o examine the  e f f e c t o f more n a t u r a l c o n d i t i o n s o f l i g h t production i n • p l a n k t o n i c  the  nutrients  systems  been  non-turbulent  study.  Experiments  culture  has  a one-stage or  r a n g i n g from  temperature 1.  dissertation  rates  during  during Experiment  outdoor  of  rearing  the importance  the  in  production  (1972), S n r a and  turbulent continuous cultures i n either  in  control  at various c o n c e n t r a t i o n s of phytoplankton.  structured to  constant  tc  by T e n o i e and  indicate  the p h y t o p l a n k t e r s , as w e l l  duration,  scale  planktonic  environment.  s t u d i e s by Walne  and  at  as  collection  trays  natural  experiments  system,  space  improve  1967),  1969,1971)  iceding  The  tc  breeding  ( 1 9 7 6 ) , and  uptake r a t e  and  the  a c o m b i n a t i o n of these f a c t o r s ,  from  including  Halne  time  of e i t h e r  p r o d u c t i o n o f p h y t o p l a n k t o n and  (Quayle,  concentrated  cr  investigations  increase ; production  bivalves,  level  o r more u s u a l l y ,  organization  bivalves  the s i g n i f i c a n c e  and  temperature  on  chains.  * The term planktonic i s used throughout this study to incorporate the fact that the experimental bivalves were s u s p e n d e d i n the w a t e r c o l u m n a n d f e d on the p h y t o p l a n k t o n . ,  8  In  Experiment  1  (Chapter  3),  duplicate  u p w e l l i n g s y s t e m s were used  t o examine  phytoplankton  the r e s u l t i n g  population  of  Experiment*^ both  scallops,  2 and  Experiment  4,  at  1.00  ).  day—*  Experiment r a t e s of  5  the  (Chapter  (Chapter  as a  include  an  system  using a  the  of  for  design  the  in  day  and  situ . ., I n  1  of  «as  primary dayr*  expanded  of v a r i a b l e experimental  two-stage  experiments,  summarized i n C h a p t e r  and 10.,  the  in  flushing period  culture  of  primary  s i m u l a t i o n model, i s p r e s e n t e d i n C h a p t e r of  and  phytoplankton  a n a l y s i s of the dynamics of the  discussion  of  discussion  (0.50  the r e s u l t i n g the  an  i n a turbulent  dynamics rates  natural  production  examined  flushing  during  source An  of  r a t e o f 0.5  examination  9 provides a comparative  five  of  the s u i t a b i l i t y food  growth  5 presents the r e s u l t s  experimental  primary  dynamics o f a  one-stage  f o o d c h a i n s was  analysis  The to  4) , the  comparative  (Chapter 7 ) .  community, Chapter  an  two  6 ) , and  communities oysters  at a f l u s h i n g  system. , C h a p t e r  communities  the  3  o y s t e r and s c a l l o p  upwelling of  community and  the  ncn-turbulent  8.,  results  of  the c o n c l u s i o n s from t h e s t u d y  are  9  CHAPTER 2.  Experimental All  EXPERIMENTAL  l a r g e tank f a c i l i t i e s  system  B.C.  production resevoir  total  a t the P a c i f i c  The  incoming  seawater  system  to  v=flow  rate  rate  (1  by  using  the  r e s e v o i r s and fittings,  nitrate  flow rate  English  Bay,  l from  *)  -  t o the  the seawater  day  - 1  )  and  by i n - l i n e  experimental  a registered  the  tanks,  trademark.  volume ( 1 ) . The  constant  during  the  method f r o m t h e r e s e v o i r ,  valves.  'inert*  a n d p l e x i g l a s s and t y g o n  * S denotes  V=tank  relatively  were  replace  ) = v V~* ;  be a t t a i n e d u s i n g  system  to  i n one d a y ; t h a t i s  a gravity-feed  the flow c o n t r o l l e d  in  in  by t h e f l o w r a t e r e q u i r e d  remained  1.0 day-* c o u l d  used  West  f o r the experimental  60 f e e t  volume o f t h e e x p e r i m e n t a l t a n k  experiments with  Institute,  t o t h e e x p e r i m e n t a l t a n k s was e x p r e s s e d as t h e f l u s h i n g  (FH), determined  flushing  using  and p r o v i d e d a s o u r c e o f c o o l ,  The  FR ( d a y * where  conducted  Environment  ( 11 PC , 20 uM  resevoir.  were  seawater  through a 5 micron f i l t e r  nut r i e n t - r r i c h  rate  experiments  was pumped f r o m a d e p t h o f c a .  passed  ANE METHODOLOGY  facilities  continuous culture  Vancouver,  FACILITIES  this  Flushing  method.  rates  up  A l l materials  , including the fibreglass 1/2  M  PVC&  samplers.  1  piping  and  10  One-stage C u l t u r e Experiment The Figure  indoor  1,  tanks./  facilities  duplicate  l i t r e s and  the depth  upwelling  r a t e o f 0.4  the  tank  holes  hy  using  every  5 cm  from the  end  of the  fts  illustrated  B)  were s e t up  3.  Each  volume)  which  diameter,  mid  In-situ  a volume o f  720  metres.  The  0.8  over  the  inflow  samplers  experimental  whole a r e a  pipe  with  were l o c a t e d  and  bottom depths at h a l f the  to  the  centrally  ca.  and  was  siphon.  the  of  small i n both  distance  located  levelling  Natural Forcing  Conditions  (0.75  sampled situ  a.,  from  a covered large  was  d u p l i c a t e tank for  Experiments  (3200 1  production  (3.0  inflow  towards  sampling  resevoir outflo*  tank  pipe  near the  the  centre  tube,  osing  were l o c a t e d a t  of blackened tygcn used as s i p h o n s .  2  seawater r e s e v o i r  as  i n f l o w i n g seawater t o the  1  systems  three  tubing  a  m  bottom of  d i r e c t e d p e r p e n d i c u l a r l y upward  to the  The  samplers  Three long pieces a PVC s t a n d p i p e and  angled  m)  the  3,  outdoor platform  t a n k was  f o r Tank  In  an  water f l o w  distance 3  on  2 and  7500 1 v o l u m e ) . . The  tank,  Figure  i n Figures  gravity-fed to a  production  the  Experiments with  c o n s i s t e d of  of the  1  constant  in  ;  (A an  tank  m/day was  tank  One-stage C u l t u r e  half  t a n k s had  Conditions  illustrated  the  water c o l u m n was  apart.  surface,  1 are  views o f  a rectangular-shaped  the  and  side  production  of the  tanks at  pipe.  f o r Experiment  i n c l u d i n g s u r f a c e and These  with. C o n s t a n t F o r c i n g  shown  the at in  production tygon-tubing  »stations* i n  were i n s e r t e d i n t o  11  the  c e n t e r of  and  bottom  m..  The  was  sampled  the  production  (1.0m).  The  tank: surface  total  seawater o u t f l o w  from t h e  using a T-joint  production  tanks  p l e x i g l a s s to  were  reduce  depth of  with  the  a reduced with  the  thin  contamination  (0.5m) was  1.1  production  tygon  a  mid  water c o l u m n  surface of  covered  aerial  (0.05m),  tank  fitting.  The  (1/8") s h e e t  and p h o t o - i n h i b i t i o n .  T w e l v e s u b s t a t i o n s were e s t a b l i s h e d i n each p r o d u c t i o n for  location  of the  illustrated the  i n Figure  growth o f  commercial per  the  The  and  horizontally  herring  experimental  at the  were  s u r f a c e , mid  E x p e r i m e n t s 2 and  frame  net.  experimental  illustrated were  were no produced Outflow  herbivores in  three  the  primary  (170 and  the  and  giqas  ,  bottom  depths  and  Chlamys  to r e c t a n g u l a r X  30cm  net  20cm  depths  in  Tank  Natural Forcing  B  for  5. . The  Conditions  E x p e r i m e n t s 4 and  primary  5  production  and A  turbulent diffusion and  B by  tank c o u l d  litres), other  be  fed i n t o  d e n s i t i e s of o y s t e r s  as  the Two  experimental  ( 16,  3 2 and  48  there  actively  using submersible  shown i n P l a t e I . four  was  are  tanks  one-stage c u l t u r e system, except  Tanks  from e i t h e r  accligation  examine  facilities  i n situ  both  h e r b i v o r e : tanks for  to  as  3. .  i n F i g u r e s 4 and  identical  3,  c a g e s were s u s p e n d e d  bottom  Two-stage C u l t u r e Experiments with The  tank  used t o e x a m i n e  ( ca.  These  or  and  herbivores,  confined  a plexiglass seine  2  Crassostrea  a t s u r f a c e , mid  .faerieia • • ( s c a l l o p s ) ,  )  during  other  with  Tank A was  oyster,  were s t r u n g  cages, constructed X4cm  during Experiment  2 f o r Tank B.  Pacific  cultch  substation.  hastata  herbivores  of  pumps. smaller  were  used  tanks  to  oysters  per  12  tank r e s p e c t i v e l y ) , plus a c o n t r o l in  situ  circulatimg  Throughout identification  pumps t o k e e p  the;  text,  (no o y s t e r s ) . the system  A  and  B  had  homogenous.  are  of experiments as a reference  A l l tanks  included to the  in  the  appropriate  t a n k s y s t e m . ,-  Methodology The  same  experiments.; •stations* mid(M), (PST)  sampling  Water s a m p l e s  procedures  were  ( 4 litres)  were, t a k e n f r o m  i n the p r o d u c t i o n  bottom(B)  and  system  outflow(O)  The  samples  experimental minimize  animals  handling.  variables  Physical During provided  A  Vitalitefi in  T h e r e was langley  which  0800 and  i n PA8  -  the and  the  five  s u r f a c e (S), 0900 h o u r s  and  biological  the  laboratory  Measurements aquarium  on  the  wet-lab to  derivation  pertained  of  t o the primary  Temperature  1, c o n t i n u o u s fluorescent  the  systems  the  in  radiation simulated  the  culture  o f e a c h t a c k was  was solar  range  (Pfifi) f r o m c a .  radiation  between the d u p l i c a t e  at the periphery  Salinity  t u b e s , which  radiation in  and  artificial  composition  available  no d i f f e r e n c e  min *  in  in  for a l l  1.,  spectral  photosynthetically  below.  description  i n Appendix  Experiment  analyzed  made  Parameters; L i g h t ,  by  radiation  decrease  were  and p a r a m e t e r s  a r e summarized  0.10  ) between  were  a c c o r d i n g t o t h e methods o u t l i n e d  ;  inflow (I),  f o r t h e measurement o f p h y s i c a l , c h e m i c a l  parameters.,  nm,  (  followed  of  400-700  intensities  of  t a n k s , and a 5% not  considered  13  a significant  reduction.  During  the  the  incident  the  plexiglass  solar radiation  mill!voltmeter  the  primary  radiation  estimates  (  Similar  r e s u l t s were  submarine  continual  problems  produced EXTK  extinction  based  which proved  •  into  Experiment  • 0.0  1,  the  plexiglass light  the  curves  period  based  that  the  of  on PAB  of  daily  Estimation  was  the  the  was  was  the  Suckling was  not  (1966).  placed  radiation  coefficient leakage  at  i n t o the  a  a  r  A the  depth  However, photocell  estimates  of  between  the  .relation  chlorophyll  in  (EXTK).,  consequently,  empirical  concentration:  (.054* (CHLA**.66667) ) chlorophyll^  minimal  measured  Szeicz  solar  seawater  Biley's  by  1966)  ( Siley,  to  seawater sample.,  in situ  recording  t h e n summed f o r ).,  - 1  found  (.0088*CHJLA) •  were  The  a  (r*=0.S6)  r e l i a b l e f o r a range of  Temperatures  1Q8.  (PflB)  measurements;  other p a r t i c u l a t e matter  inserted  SB  extinction  c o e f f i c i e n t and  EXTK = .04  day  (Haddux,  with  on  by  The  under  using  YSIS  of  and:  0.50  the  tanks,  hour).  radiation  determine  unreliable  were  per  previously  e x p e r i m e n t a l tank t o measuring  2"  recorded  calibrated  experimentally  photometer  (PARZ) by  a  radiation  cover, =  experimental  with  langley  active  PAB  measurement  continually  e x p e r i m e n t s , and  determined  of c l o u d  was  outdoors,  hour i n t e r v a l s , i n c l u d i n g  productivity  who  function  solar  for four  photosynthetically (1974)  speed=  incident  were i n t e g r a t e d  the  precision)  (chart  the  (SB)  sheet covering  (5%  solarimeter  reduced  experiments conducted  thermistors  0.1  .PC  During  values  when  1956)• using a  the  connected  thermometer  first to  a  half  of  2-channel  14  Bustrakfi  , recorded  the  p r o d u c t i v i t y tanks,  two  thermal  temperatures  instability.  occurred  i n the tanks  at the  surface  and  p r i m a r i l y t o d e t e c t any  Temperature  fluctuaticns  only during  the t e n  of  bottom  of  evidence  of  ca. •  0.2  minute sampling  °C  period  e a c h day.,. Salinity from  the  samples  i n f l o w and  Nutrient  were  Strickland  and  determined  Technicons  (1972). using  in  i n the  days of c o l l e c t i o n ;  ho*ever>  days.  The  outlined  Daily  samples  for  the  range  of  in situ  d i f f e r e n c e s between t h e significant  cadaium-copper  usually  there  p r e c i s i o n of t h e  was  concentrations  Ammonia  i n e i t h e r case  phenol-hypochlorite contamination  of  using  method  0.26  The  uM  within  methods  by  acid  uM  N l  -  care  of  over )  _ 1  were  at t h e f i r s t  a  change  duration  N 1  a  samples  measured  ( 0 - 25  automated  Although  double deionized  ( S.E.=  o r . with  sign  and not of  , 1972).,  were d e t e r m i n e d m a n u a l l y u s i n g  method. samples  nitrate  significant  2%,  in  reduction  analyzed  was  (Hager e t a l .  concentrations  5..  a storage  s i n c e t h e c o l u m n s were r e p l a c e d  deterioration  1 t o 3,  no  with  method  manual and  silicate  methods  E x p e r i m e n t s 4 and  d a r k and  and  the  i n conceutrations of n i t r a t e  f l a s k s and  bottles  salinometer.  manually i n Experiments  Auto-analyzer  were s t o r e d a t 2 °C  of t h e  using  were a n a l y z e d  c o l u m n method, e i t h e r  the  u s i n g ia EeckmanS  analyzed  Parsons  determinations  eight  in salinity  ammonia, u r e a , r e a c t i v e p h o s p h o r o u s  nutrients  (P<»01)  daily  Parameters  Nitrate,  few  were c o l l e c t e d  was  taken  to  c l e a n i n g glassware,  the  reduce covering  distilled  water, t h e p r e c i s i o n  ; n=3  poor r e l a t i v e  1  ) was  to  15  the  low i n s i t u  analysis  was  problems of samples  concentrations based  the  on  the  ammonia  were  (  <  2.0  urease  for  a  few  increased  Degobbis,  1973)..  p h o s p h o r o u s and s i l i c a t e be  the  like  primary  many c o a s t a l  Primary  urea  analysis,  n u t r i e n t s may  Only o c c a s i o n a l  limiting  Urea  and  before  have  measurements o f  were t a k e n s i n c e n i t r o g e n  nutrient  ).  1  ammonia  weeks  i n the c o n c e n t r a t i o n s of these  1~  with the i n h e r e n t  Since  variabilities <  N  method  analysis..  frozen  uH  appeared  phytoplankton  to  productivity,  ecosystems.,  Parameters:  Standing Stocks Primary  P r o d u c t i v i t y , and  Community S t r u c t u r e The every  standing stock of phytoplankton  day  at a l l four  chlorophyll  a  (CHLA) , a s  (1972)..,  water  HilliporeS  AA f i l t e r s  10.0  ml  stored  of  acta  116  and  phaeopigments  spectrophotometer.  equations, although t h e SC01/UNESC0  average  Strickland  filtered  and  through  Parsons  0.8  micron  grade  based  were  acetone.  days  The t u b e s  before  determined  Calculations  on t h e P a r s o n s  were  chlorophyll,  using a of  tetween  Beckman pigment  and S t r i c k l a n d  t h e r e was no d i f f e r e n c e  in  (P.S.).  these  and  (S/U) e q u a t i o n s f o r CHLA:  CHLA(P.S.) precision  in  f o r two t o s e v e n  were  measured  and t h e p i g m e n t s e x t r a c t e d i m m e d i a t e l y  spectrophotometric  concentrations  The  were  usually  s t a t i o n s a s the c o n c e n t r a t i o n o f  outlined  samples  i n a freezer  carotenoid  in situ  was  = 1.0 * CHLA (S/U)  '<r*=1..0; tt=41)  o f t h e method a t t h e 30 ug C h i a 1-* l e v e l  standard e r r o r  o f 0.5 ug C h i a  duplicate determinations.  l  -  * ,  based  on  was an eight  16  To  estimate  the  plates  were s u s p e n d e d  x 6 cm  glass slide  from  both  filter,  i n the  standing  primary  a t t a c h e d by  sides  and  phytcbenthic  tanks,  plastic  of the s l i d e  ( 20  duplicate chlorophyll  cm  stock,  plexiglass  each w i t h f o u r 2  clips,  algae  was  cm  scraped  ) cntc a preweighed  2  a and  biomass  estimates  GFC were  d e t e r m i n e d . ., Primary  productivity  t h r e e days as method  net c a r b o n  outlined  w a t e r sample  was  ampoule  5  cf  solution)  added  experimental  the  this.„ bottle  using  1300  containing  filters,  twice,  LSCS  The  .  calculate method. duplicate  the  rate  Precision  of  carbon  estimates  b o t t l e s ranged  values <  l-»  hr-*  10 ag C 1~*  Exudation  frcm a  of organic carbon  a modification of the  Zeutschel  in  (1970).  5  ml  and  for  0.45  carbon *as  micron  of  methodology of f i l t r a t e  from  the  were  required  based  on  once  pH  between of  11%  75  ug  a  week  i n Anderson »*C  to  the  variation  determined outlined  vials  Tri^Carb  for p r o d u c t i v i t i e s ^  was  HA  Samples  prcductivity  coefficient  for  After incubation,  using a Packard  **C  the  incubated  fluid.,  fixation  a  in  scintillation  carbonate  h r ~ * t o 9%  NaCl  bottle  through  placed  of  ( i n 3%  and  f o r ten minutes,  determination  for  using  each  one  holders, (PST).  ml  and  horizontally  hours  the  A 100  bottle  bicarbonate  15 ml o f A g u a s o l B s c i n t i l l a t i o n  counted  (1972).  ECD  every  using  suspended  filtered  then  hour  production  plexiglass  s a m p l e s were i m m e d i a t e l y  ml)  sodium  This  per  Parsons  (112  were  c a . , 0900 -  approximately  litre and  radioactive to  tank  MlliporeS  per  Strickland  dark  hours from  fixed  estimated  placed i n a small uC  corresponding  4.0  in  was  C  and  productivity  17  samples then  were a c i d i f i e d  bubbled  inorganic  with  vials  fluid  prepared  Oxygen bottles, and  containing  samples  in  were  the  of  was  significant  bottles  variance  stored  respiration  BOD  this  during  of  the  using  coefficient level,  of  1.0%  at  oxygen I *  of  period  as  the  300  ml; s u b s a m p l e cc  mg  the  method  based  0.831  oxygen  (250  species was  ml)  immediately  day  light  or  two.  that  1  An there  between and  dark  bottle  Light by  1.2  and  a  carbon  respiratory  to carbon  duplicates  the  325  mg  hr-*  and  7.5%  and  plexiglass  radioactive  conversion  at l "  and  1972).  on  BOD  units. was  oxygen  l ~  at the  a hr~  l  250  l  mg  were a l s o c o l l e c t e d s e m i - w e e k l y  c o m p o s i t i o n aDd  sample  was  stored  be  counted  on  a Model  and  Parsons  ( 1 S 6 7 ) . , The  at  2  B coulter  Zeiss °C  for  size determination. , a  f i x e d with l u g o l s o l u t i o n  s e d i m e n t a t i o n chamber w i t h a  remaining  ml  productivity  the of  to  level..  Water s a m p l e s phytoplankton  used i n  v a r i a t i o n of  hr-*  -  the  a  horizontally  1.0  precision  i n 300  indicated  Parsons,  quotient The  any  scintillation  fixed  within  the  a photosynthetic quotient were  remove  minutes.  were  experiments., of  10  Primary  suspended  same  and  i n oxygen c o n c e n t r a t i o n  manner.  were  3.0  were t r a n s f e r r e d  a daily basis  3  of  minutes to  foe  Experiment  estimated  pH  aguascie  They  ( S t r i c k l a n d and  bottles  holders  ml  1.  difference  were a l s o  oxygen t e c h n i q u e  10  20  to a  aliquots  t a k e n on  during  in  for  dark f o r a n a l y s i s  analysis no  acid  samples c o u n t e d  except i n Experiment  stored  dark  gas  carbon.,, 2 ml  scintillation the  phosphoric  nitrogen  radioactive  and  with  and  examined  inverted  i n the  Counters  a n a l y s i s employed  dark as a  in  a  microscope. , until  outlined  i t in  10 5 The  could Sheldon  flow-through  rate  18  of 0.044 ml s e c — i  using  a 100 m i c r o n  diameters  ranging  from  triplicate  and then  averaged.  Secondary  Scallops  before  to  { Chlamys h a s t a t a  by d i v e r s  the  the  seawater of  start  from 50  weighed 45.  1 (EXP1). . T h e y  Experiment  tank  Bay were added scallops  adhaerens  tank  The l a r g e r rejected. 1  litres  t o t h e tanks  were s e l e c t e d  ).  seawater  of surface  every  couple  as experimental  sponges  ( Myxilla  The s c a l l o p s were  t o t h e benthos  weeks  tagged,  o f Tank A on and  Day  weighed  growth.,  2A  Crassostrea gigas cultch  holding  Ten  two  of  one month, t h e C h l a m y s were r e - m e a s u r e d  Experiment  near  1).  at ca.  were p u t i n t o  supply  were c l e a n e d o f e n c r u s t i n g  fiycale  to deterBine their  grower  Columbia  a constant  (Figure  and m e a s u r e d , and added  After  ) were c o l l e c t e d  of  healthy  ,  bericia  British  English  a n i m a l s , * and a few incrustans  were c o u n t e d i n  near V i c t o r i a ,  resevoir  days.  28.5 m i c r o n s  particle  1  h o l d i n g . t a n k s and p r o v i d e d w i t h from  Eleven  Productivity  Experiment  20 m e t r e s  2.82  aperature.  Victoria  on  with u a f i l t e r e d scallops  were o b t a i n e d  April  25  and  from  commercial  k e p t i n a 3000  running seawater  with the b o r i n g  a  ( ca.  litre  29 p p t ,  sponge C l i o n a c e l a t a  13 were  19  °C  ).  10  holding for  litres  of  tank e v e r y c o u p l e o f d a y s .  identification  smaller  weight  in  measured b e f o r e culture  their  experiment  at  grown f o r f i v e the  addition  on Day 6.  substations  growth  cultch  from  Tank  1  to  the  to  t o the tagged  bottom  4 (Figure  weeks b e f o r e t h e f i n a l  The  total  per c u l t c h  one-stage  Four s t r i n g s , and  were  A.  and number o f o y s t e r s  ( 0 . 1 m ) , mid (0.5 m)  located  were added  a t t h e b e g i n n i n g o f EXP2ft t o  the o u t f l o w  water  stock  Twelve  and t r a n s f e r r e d  tanks r e c e i v i n g  weight,  surface  phytoplankton  continuous  with c u l t c h  i(0-9m)  were  a t the  depths,  2 ) . . The o y s t e r s  were were  measurements were made  of  variables.  E x p e r i m e n t 2B At  the  collected  beginning  of  a t t h e same l o c a t i o n  as e x p e r i m e n t a l animals. into  They  f o u r groups o f i n c r e a s i n g  scallops, located at  EXP2B,  one-half  were  t h e bottom  station.  larger size  days  in  After  the scallops  small  transferred  on  and 64 were  size,  selected  The s c a l l o p s tanks  were weighed  were  selected  and w i t h i n e a c h g r o u p o f 16 f o r a cage  t o be  f o r a c a g e t o be l o c a t e d  The same p r o c e d u r e was a d o p t e d  groups. covered  scallops  were o r d e r e d ±y l e n g t h a n d w e i g h t  randomly  were a c c l i m a t e d  f o r the f o r four  f e d by t h e o u t f l o w f r o m Tank B. and  measured,  Day 10 o f EXP2B t o t h e i r  The f o u r c a g e s a t t h e mid d e p t h 4,  i n Victoria  a t mid d e p t h a n d t h e o t h e r h a l f  three  to  s i x dozen  s u b s t a t i o n * s 5,8,9 and 12 ( F i g u r e  2 ) • One  measurements were made o f t h e g r o w t h  r  cages  were  designated substation.  were l o c a t e d  w h i l e t h e f o u r c a g e s a t t h e bottom  the  at  depth month  variables. ,  substation*s  1  were l o c a t e d a t later,  final  20  Experiment The  24  transferred outflow  3A  oyster  cultch  d u r i n g EXP2A  f r o m Tank A.  Pinal  depths,  The  were  Another Victoria EXP2B  was  this  four  nine dozen  scallops  Day  adopted  7 o f EXP3B.  for  the  (EXP3B),  (containing  8 scallops  acclimated  were  t h e y were w e i g h e d 11 t o t h e  The  Juvenile  as w e l l  and  tank  constantly  A.  at  the  same  of i n c r e a s i n g  per cage)  size.  to  string  a t the s u r f a c e  as t h e mid  and b o t t o m  depth  stations.  f o r t h r e e d a j s as i n EXP2B, and  SUBSTN  in  were  Tank - B.  variables  after  transferred The one  final month.  5 Crassostrea  until  Unfiltered  supplied  g i q a s were o b t a i n e d from a  Victoria, British  outdoors  experiments.  and  o f the 96 e x p e r i m e n t a l  measured, the c a g e s  appropriate  o y s t e r grower near litre  mid  same p r o c e d u r e used i n  classes  measurements s e r e made o f t h e g r o w t h  Experiment  Day  growth.  t h e r e " e r e enough s c a l l o p s  The  Day  the  1 t o 8 i n Tank  collected  selection  11)  after  surface,  weeks o f  were  g r o u p i n g them i n t o f o u r  scallops  by  3B  experiment  cages  two  and  were measured on  at s u b s t a t i o n s  ( S U B S T W s 6,7,10 and  on  variables  tanks fed  w i t h c u l t c h a t the  located  l o c a t i o n on  a n i m a l s and In  growth  measurements were t a k e n a f t e r  Experiment  f o r EXP3A were t a g g e d  to the s n a i l h o l d i n g  6 o f EXP3A, a n d 8 s t r i n g s bottom  selected  to  the  running the  Columbia start seawater  holding  tank  commercial  and h e l d i n a 3000  cf  v  the  hervivore  (29 p p t ,  13 °C)  was  and  litres  of  10  21  phytoplankton A  small  wiring  hole  cultch  s i z e groups  Details experiments (length,  a s growth  (Plate could  are  of  days.  i n t h e unto o f the s h e l l f o r  square p l e x i g l a s s boxes d e s i g n e d  I I ) , so growth  variability  between  to and  be a s s e s s e d .  presented i n Chapter  7.  continuous 1 be l i n e a r  depth) , w e i g h t i n a i r ( i 0.1  g),  culture  dimensions weight  in  measured  variables. and  t o t h e h e r b i v o r e growth •  drilled  whole volume o f t h e i n d i v i d u a l o y s t e r s were  A description  .  t o the t a n k e v e r y C o u p l e  of the d e s i g n of the two-stage  w i d t h and  w a t e r , and  2.  was  (1/32**)  the o y s t e r s t o small  simulate within  s t o c k were added  the d e r i v a t i o n experiments  of the  variables  a r e summarized  in  pertinent Appendix  22  CHAPTEB 3.  ONE—STAGE GQNTINOQOS CULTOBES IK  NON-TUBBOLENT  OPWELLING SYSTEMS BITH CONSTANT FOBCING CONDITIONS  The examine  initial  p h y s i c a l and  turbulent The  experiment  biological  examination  of  the  constant, forcing conducted  for  samples  ten  phytoplankton  phytoplankton  weeks,  each  stock  English  Bay  and  screen  to  remove  to  determine  for  the  passed any  in  situ  been  flushing  Tank  problem  A  levelling on  a  small  zooplankton.  under  controlled,  1.,  of this  and  the  initial  litres  of  the s u r f a c e o f  diameter  (54u)  wire  tenthic herbivores, Day  continuous c u l t u r e  45  system  scallop population.  Communities  relatively  although  as 0i45 d a y * -  i n c r e a s e d by  .  on  a  c o n s t a n t a t 0.5 few  days  However cn Day  2 i n c h e s due  31  overflow  o c c u r r e d i n Tank B o r Day  pipe 43  since  although  day  1  the  rate  the  water  to c l o g g i n g of  p i p e by a f i l a m e n t o u s N a v i c u l a mat the  was  After  10  frcm  .  - 1  the  (EXP1)  with  The  day  , were i n t r o d u c e d i n t o Tank A on  r a t e remained  t o a s low  seeded  0.5  non-  permitted  seawater  been c o l l e c t e d  through  the experiment,  growing  was  2,  on O c t o b e r  inflowing  to  duplicate  experiment  growth o f a l o c a l  Dynamics o f t h e P r i m a r y  in  tank  the s u i t a b i l i t y  s u r v i v a l and  throughout  This  conducted  rate of  dynamics  beginning  which had  Chlamys h a s t a t a h e r i c i a  level  in  described i n Chapter  with f i l t e r e d  taken,  decreased  and  variables  conditions.  were f i l l e d  The  designed  u p w e l l i n g systems with a f l u s h i n g  experimental system,  tanks  was  the  which  had  The  same  the s i t u a t i o n  was  Day  13.  (  23  more  severe  since  t h e t a n k o v e r f l o w e d f o r about h a l f  In each c a s e , the N a v i c u l a around  Physical The  flux  of  were  of  144  0.10  weeks  langley  day  the 12.0  °C  °C  Nutrient The  °C  essentially first  (± -1 and  °C  10.5  two  eight  ) a t the °C  (± . 1  weeks,  the  3 °C a t t h e s u r f a c e o f t h e and  bottom  of the i n f l o w i n g ppt  the  last  (± 0.6  stations.  seawater  was  ppt) d u r i n g  The  constant EXP1.  start  N  l - i ,  71.3  ufl  concentrations  of  the  nutrients  sere  measured a t v a r i o u s t i m e s d u r i n g  Samples of the i n f l o w i n g of  concentrations: nitrate  (SI03) =  ca.  temperatures  but  station  the  mid  oc ) and 2.9.2  productivity  the  UH  the  concentrations  the experiment.  0.88  for  radiation  Conditions  phytoplankton  before  °C a t  depths  depths  During  decreased  and s a l i n i t y <± .1  between  in situ  a v e r a g i n g 14.5  station.  ;  10.0  resulting  within  available  represented a daily  (± . 1 °C ) a t t h e mid  t a n k s a n d c a . , 1,5  at  The  different  temperature  temperature  photosynthetically  ain-*  experiment,  °C ) a t t h e bottom average  t h e p i p e were removed.  of  .  - 1  between t a n k s and  of  surface,  source langley  significantly  constant  hour.  Environment  continuous  radiation  an  EXP 1,  (N03)  =  phosphate Si  1~*  suggested  . that  with  17.8  uH  <P04) = The the  important  in  s e a w a t e r were a n a l y z e d the  resulting  average  H . I r * , ammonia  (NH3)  1.75  silicate  ratios  ufi P 1~* of  these  experimental  ,  =  nutrient  system  would  24  probably  be n i t r o g e n - H a l t e d  The  concentration  Figures  6  averaging first in  23.6  five  situ  first  and  of n i t r a t e  7.The uH  (SE = 0.39  The  pattern  nitrate concentration  three  recovery  weeks, w i t h  to  levels  during  high i n f l o w  N 3>»  weeks.  * .  an  of  NG3 .ca.  15  N03  fifth  as  high  correlation  B,  although  The  as  N 1~  at  the  The  this  may  have  Day  4 3.  on  average  weekly  uH  the  bottom s t a t i o n the s u r f a c e  and  n e v e r , l e s s t h a n 0.5 1-*  ,  As  greater  P I  indicating  by Day uM  N l~  and  -  1  i n the c a s e  P04  were  a  high  to  of  about  negative for  the  tank  was  P04  at  concentration  Phosphate  not  the  phosphate  c f N03,  phosphate  did  and  stations  due  , e v e n when N03  that  B  fourth  c o t a p p a r e n t i n Tank  partly  stations.  Throughout  the  bottom  was  then a  significantly  Tank was  Day  and  .,  l  During  in  than the  15,  was  inflow concentration  outflow uH  been  the  10,  There  surface  for  U H N I T * -by  station  same t r e n d  P I-* to b o t h t a n k s . was  A.,  the  the  1  o f 2-5  botton  Tank  during  between t a n k s f o r  concentration.  o f EXP1 A.„  2.43  uB : N  in  between N03  duration  overflowing  at  a t the  surface  ;n=35)  in  constant,  damped o s c i l l a t i o n s  weeks, t h e a v e r a g e n i t r a t e . l e v e l s  twice  the:  the  1~*  uM  the  than  u«J  relatively  minimum o f <2  concentrations  greater  was  similar  decrease to c i t r a t e experiment, the  BC3  of  was  EXP 1 i s i l l u s t r a t e d  values  levels  were  were c a .  limit  2  primary  productivity. After  the  introduction  d i f f e r e n c e between t h e  net  of  the  s c a l l o p s to the  concentration  of  system,  ammonia  at  the the  R e s u l t s from A n t i a e t a l . (1963) indicated that a similar primary community had a N/P r a t i o g r e a t e r t h a n 12, w h e r e a s the N/P r a t i o o f t h e s e a w a t e r s o u r c e was o n l y .10. 1  25  bottom s t a t i o n  was  )  (0,10  t h a n Tank E  Phytoplankton  Standing The  uM  N 1~»  of  ).  During  the  this  initial  at  EXP1.  The  on  two  of the f i v e  Days seeks  magnitude  n o t a s l a r g e i n Tank  of the blocm  benthic  B.  54.5  system  which  was  produced  averages  a l - i (SD=6.49) 16.1 l-i  ug  of  during 18.8  the  ug C h i a h  a  and  both  reduced a  third  ( F i g u r e s 8 and depth  9).  throughout tanks,  p h y t c p l a n k t o n maximum  sp., the  , PAB  the  fact  which  was  more  at  depth  A comparison  post-blccm 1  period  ug C h i a 1-*  in  (t>t0), ug  (SD=9.17)  at  29.1 the  a  of CHLft  (SD=5.19) and 7.4  ug C h i a l - i <SD=16.?4)  was  from  C h i a - l - i (SD=9.88) a t t h e n i d d e p t h , and 27.Q  9,  a  ug C h i a l ~ i i n T a n k s A and  Navjcula  a t t h e s u r f a c e , 21.2  (SD= 18.57) and  depth..  1  stock  in  t h e same i n b o t h  already l i g h t - l i m i t e d . B  later  with  was  f u r t h e r reduced  between t a n k s A and  and  This probably resulted  diatom,  p r e v a l e n t i n Tank B,  8  week p e r i o d  B r e s p e c t i v e l y , although the secondary  the  -  five  phytoplankton  oscillations  increased significantly  a v e r a g i n g 5 2 * 9 ug C h i a 1-* and  that  the  damped  bloom  increase  stock levels  time,  of  maximum a p p r o x i m a t e l y t h e end  measured a s t h e c h l o r o p h y l l  t o t h r e e days f o r the f i r s t  secondary  The  N I  y  s t o c k was  (CHLA) e v e r y two  EXP 1..  with  uM  Stock  generally followed a pattern tanks,  h i g h e r i n Tank ft (1.50  Dynamics  phytoplankton  concentration weeks  significantly  Chi and  ug C h i a bottom  26  After  the  phytoplankton the  bottom  stations  scallops  s t o c k was r e d u c e d station,  and c a .  both  benthos as  tanks .  1  which  added to less  was  of  the  The p h y t o b e n t h o s  times the  phytobenthos  an  average  -  1  than  the  corresponding  b l o o m , e s t i m a t e s were  made  s t o c k which sank t o t h e  was measured  cn Days 9, 10 a n d 14 (±  11  (± 11 ug C h i a 1~* ) , which -  1  day  ....  - 1  c o n c e n t r a t i o n was a p p r o x i m a t e l y  concentration net  at  t h e s u r f a c e a n d mid  v a l u e o f 23.4 ug C h i a I  phytoplankton  representing  l  (± 5 ug C h i a 1-* ) , 256 ug C h i a 1-*  an average  the  than  phytoplankton  ug , C h i a l-» ) and 343 ug C h i a l - i  average,  t h a n 5 ug C h i a  less  During the i n i t i a l  180 u g C h i a 1~»  represented  t o Tank A on Day 45, t h e  40 ug C h i a 1~» l e s s  v a l u e f o r T a n k B. in  were  sinking  in rate  the  water  On 2.5  column,  o f 1.0 m/day o r 1.4  m/day w i t h r e s p e c t t o t h e water c o l u m n .  ?  Primary  Productivity  Primary  productivity  r a t e s c n h o u r l y b a s i s were low i n b o t h  tanks d u r i n g the experiments less  than  30  ug  C  (Figures  10 and 1 1 ) .  (ASS)  the bottom s t a t i o n Day  19  similar  as  were  1~* h r * , e x c e p t on Day 49 a t t h e bottom -  s t a t i o n . . . When s t a n d a r d i z e d on a s t a n d i n g s t o c k productivities  Values  ranged  from  0.0 ug C (ug Chla)-»  t o a v a l u e o f 5.1 ug C  illustrated  in  basis,  (ug  Chla)-  1  primary hr-* at hr  -  1  on  F i g u r e 12., The v a l u e s o f ASS were  between t a n k s b u t d e c r e a s e d  significantly  with  depth.  * C i r c u l a r p l e x i g l a s s c o l l e c t o r s were c o n s t r u c t e d a n d p l a c e d a t known locations on t h e bottom of the tank b e f o r e the s t a r t o f E X P L y T h e s e c o l l e c t o r s were r e t r i e v e d by p l a c i n g a covering plate (with an 0 - r i n g o n t h e i n s i d e s u r f a c e ) a t t a c h e d t o a n i n f l e x i b l e 1 metre r o d , o v e r t h e b e n t h o s c o l l e c t o r a n d bringing i t to t h e s u r f a c e .  27  Excluding  Day  19,  the  experiment averaged ug  C  (ug Chla)-»  (av  SD=.26)  at  standardized  0.96  hr~4  ug  C  productivity rates during  (ug C h l a ) - »  (av SD=.21) and  the  surface^  hr~  0.40  mid  (av SD=.40),  l  ug C  and  th 0.76  (ug C h l a ) - * h r - i bottom  stations  respectively.,  Phytoplankton Stock During  the  Composition  initial  bloom,  dominant ; p h y t o p l a n k t e r , diatoms  (  station sixth  of  were a l s o p r e s e n t .  the  third  ,  other  and  fourth was  weeks. , By  not  present  SP.  Havicuia  ),  sg.  a  green  were  was  replaced  and  B h i z o s o l e n i a sp.  A.  Chaetoceros  sp.  by  became t h e  b o t t o m s t a t i o n compared  F l a g e l l a t e s i n t h e 2-20 T a k a h a s h i (1973a), p . 6 ) .  with  at  the  sp.  numbers  end  of  the  surface  , Gymnodinium and  the  phytoplankters. mid  and  s c a l l o p s were added dominant  size  surface  i n the  sp.  and  bottom  , T h a l a s s i o s i r a sp.  range  to  Phytoplankter  T h a l a s s i o s i r a SP.  micron  , Nayicula  the  dominant  H i t z s c h i a spp. After the  of  flagellate,  the  the  species  Increased  Three d i n o f l a g e l l a t e s ( p e r i d i n i u m Dinophysis  was  were a p p a r e n t a t t h e  dominance of Skeletonema costatum  stations  1  *  week. S k e l e t o n e m a c o s t a t u m  filamentous The  )  nano-flagellates  during  samples. s£.  although : several  costatum  N J t z s c h i a spp. , , T h a l a s s i o s i r a scp.,  # Chaetoceros sp. diversity  Skeletonema  i n Tank  (See  ,  Tank at  the  B.  Parsons  and  28  Growth and  The  Survival  results  turbulent day-*  )  of t h e S c a l l o p  of  Experiment  upwelling  provided  system  a  suitable  after  the  a  on Day  phytoplankton were  often  indicating The weighed the  Tank  a t t h e bottom observed  50 5.Og  station  in  t h e 5.0-5.9 cm  initially..  (SD=.29) i n l e n g t h .  weight  cm per  (av  the  the  ventral  to and  scallops  direction*  cm  and  The f r e q u e n c y d i s t r i b u t i o n  of  b a s e d on  from  3.0  length  4.0-4. 9 cm  (av 1=  at  cm  t o 6,6  (L) i n d i c a t e d  cm  range  t (0)  The  averaged  (SD=3.35) and 37.Og  7 were longer  corresponding 6.2g  (SD=. 33) ,  (SD=1C28)  for  4.5  averaged  5.4cm; SD=.22) and 5 were  SD=.33).  that  range, a v e r a g i n g  the four  week growth  experiment,  total 12.6g  the  four  most o f t h e C h l a m y s  had l a i d  down a t h i n , d a r k l y p i g m e n t e d  band  o f new  average  increase  in  length  (NETL)  and  width  scallop  population  was  0.09  (SD=. 096)  represented  d u r i n g t h e month. of  of  groups. after  which  growth  C f t h e r e m a i n i n g 12 s c a l l o p s ,  6.2cm;  scallop  (SD=2.77) , 24.5g size  range  1=  (0.5  were i n t r o d u c e d  r e d u c e d and  8 s c a l l o p s i n the 3.0-3.9  CB  rate  ;  population  3.5  the  non-  f o r r e m o v a l o f p s e u d o f e c e s .«  t o 51.7g  (SD*. 2 5 ) . , The  t h a n 6.0  was  in  30 o f t h e 50 C h l a m y s were i n t h e cm  for  the  45, t h e amount o f d e t r i t u s  s c a l l o p s ^ ranged i n l e n g t h  experimental  that  exchange  the s c a l l o p s  swimaing  t h e i r need  indicated  environment  .  of  1  w i t h a moderate  Chlamys h a s t a t a h e r i c i a benthos  Population  the scallops,  growth  ca  rates  o f 2,0%  However, NETL was and  t h e net l e n g t h  also  and  0.05  and  shell. (NETH) o f cm  1..2E  a function  as a p e r c e n t  The the  (SD=.085), respectively of t h e  increase  size from  29  the  initial  group  length  from  5.1.1 f o r t h e s m a l l  t o 2.0$, 1.8% a n d 0.0% f o r t h e t h r e e  respectively. 2.1%,  The a v e r a g e i n c r e a s e  6.9%,  larger  in total  a l t h o u g h .NETHT was a l s o a f u n c t i o n  scallops.  size  (PEEL) , r a n g e d  of  weight the  T h e s m a l l e s t c h l a a y s had t h e h i g h e s t  compared  w i t h r a t e s o f 2. 9S,  size  size  classes  (NETWT) was  size  of  the  growth r a t e s o f  1.3% .and 2.7% f o r t h e . l a r g e r  classes. The  similar  60*  depth  t o t h e depth  of the seawater i n t a k e from  c o l l e c t e d , suggesting that temperature  and  conditions.,.  higher  the  bothered  by t h i s ,  phytobenthos  the  was  probably  Although  observed  occasionally  difference  scallop  population  was was  endemic p h y s i c a l e n v i r o n m e n t o f  the  n a t u r a l environment, a n d were  and  most s i g n i f i c a n t  their  salinity  experimental than  which  at the I n s t i t u t e  fairly  similar  t o the  light  intensity  was  t h e Chlamys d i d n o t seem  actively  swimming  feeding  around  between t h e n a t u r a l  on  the  the tank.  The  and e x p e r i m e n t a l  e n v i r o n m e n t s was t h e p h y t o b e n t h i c c o n c e n t r a t i o n , and t h e r e s u l t s of t h i s  experiment i n d i c a t e d  enhanced secondary  that  productivity  this  type of upwelling  i n the s c a l l o p  system  food c h a i n . ,  30  CHAPTER 4.„ ONE-STAGE CON1INOOOS CULTURES I J NON-TURBOLENT UPWELLING  SYSTEMS «ITH NATURAL FORCING CCNDTIONS  Turbulent examined  as  growth  herbivores. forcing of  u p a e l l i n g systems a t various  To  environments  evaluate  for  the c u l t u r e  experiments  (designated  flushing In  t h e growth  0.25day—  in  promoted  2,  of  two  different  In  EXP2,  in  Tank  approximation  productivity  survival  and  Based i n p a r t  the  culture  following 1.  of  to  the  compare  and  this  communities  populations  were  at f l u s h i n g r a t e s of flushing  (  and  growth  Chlamys  of  fcastata  factors  limiting  changes i n the primary  rate  had  growth  oysters  (  hericia  )  system  a  secondary community  populations.  on t h e r e s u l t s o f EXP2, a was r e - e x a m i n e d  similar  one-stage  i n Experiment 3 with  modifications:  Tank A was s t o c k e d  with t w i c e  oyster cultch at eight substations flushing  low  A and Tank B r e s p e c t i v e l y , t o o b t a i n  caused by d i f f e r e n t herbivore  continuous  at  potentially suitable for the  q j g a s ) and s c a l l o p s  were, e x a m i n e d first  sets  mixed d i a t o m / p h y t o f l a g e l l a t e .community and r e s u l t e d  oysters.  Crassostrea  herbivore  experiments,  i n environmental conditions of  systems  the dynamics o f t h e primary  In e a r l i e r  a  two  a s E x p e r i m e n t s 2 and 3)  one-stage continuous c u l t u r e s  .  1  natural  rates.  Experiment  examined  were  planktonic  s y s t e m u n d e r more  were c o n d u c t e d o u t d o o r s i n d u p l i c a t e t a n k -  and  sessile  the  c o n d t i o n s o f s o l a r r a d i a t i o n and t e m p e r a t u r e ,  controlled  high  flushing rates  rate  (0.25  day  - 1  the d e n s i t y )  ) since  while optimal  of h e r b i v o r e s using  the  (  same  t e m p e r a t u r e s and  31  suitable  p h y t o p l a n k t o n s t o c k s were p r o d u c e d  2, Tank B was  s t o c k e d with s c a l l o p s  bottom  depths but the f l u s h i n g  to  day-* t o  0,75  these  rate  primarily  system,  at the s u r f a c e , was  promote a s u i t a b l e  herbivores  in this  increased  growth  by  mid  and  three-fold  environment  lowering  the  for  in  situ  Flushing  Rate  temperature.  Comparison  Both seawater  o f Two  p r o d u c t i o n t a n k s A and on  June  and t h e . i n i t i a l 090,0 and  hr  costatum  through  a 54u  Navicula remained  , and  relatively  experiment.  * variation  diatom  Thalassiosira  sp.  (CV)  to  phytoplankton  of  a few  the  with  E n g l i s h Bay  was  remove  f o r the f l u s h i n g  rate  t h e 5%  p i p e , reducing the flow r a t e  apparent  until  Day  The  total  duration  and  5 weeks i n Tank  33  a t which  o f Experiment B  (EXP2B).,  ,  2 was  mat  ,  rates during  coefficient  i n t o the tank.  time t h e a l g a l  The  spp.  flushing  p r i m a r i l y due  o f t h e f i l a m e n t o u s i»enthic d i a t o m , N a v i c u l a •§•£.., inflow  Skeletonema  day-* i n b o t h t a n k s  was  20  community  Nitzschia  n a n o - f l a g e l l a t e s . , The  tank,  at  were added t o the  h r . , T h i s seed  as  - 1  collected  zooplankton,,  community  , as w e l l  oyster  day  monitored  c h a i n s , predominantly  sp.  filtered  conditions  a t 1100  c o n s t a n t a t 0.25 In  filled  r a t e s a d j u s t e d t o 0,25  from  netting  p r o d u c t i o n tank  mainly and  were  the f l u s h i n g  of t h i s n a t u r a l  consisted  B  S u r f a c e seawater  s u r f a c e o f each  the  18,  C h a i n s at- a Low  p h y s i c a l and c h e m i c a l  (PST).  filtered  litres  Herbivorous Food  of  t o growth  around  the  T h i s was  not  was  6 weeks i n Tank A  removed,„ {EXP2A)  Crassostrea gigas population  32  was  added t o  were  Tank & on  strung  outlined  at  the  i n Chapter  Dynamics o f the The  illustrated both  the  Primary of  variables  forcing  to  Shere  grazing  the  periods  the  and  are  chemical  Tank  410  solar  2 and a  included  ox  the  primary  Tank  are  statistics  for  of t h e  primary  and  EXP28  i n t o the  pre-  measurements. ,  tanks  e x p e r i m e n t s , from  13)  day-* f o r an 1  .  The  110  radiation  level.,  There  the  (TEMP) f l u c t u a t e d i n r e s p o n s e t c t h e The  thermal  decrease  between  ideally  to the  two  temperatures  i n SB the  TEMP  an  significant  °C  in s i t u  on  during  b a s e d on  no  10.9  in  day-*  l a c k o f an  variability  structure  including a significant  of  ranged  period  o f 35%  was  period,  (SB)  langley  extended  l a r g e CV  experimental  similar,  as  B  values  l a n g l e y day-* i n d i c a t e d t h e  during  15).  and  f o r the  tanks  and  10,  and  breakdown  i n f l o w temperature  14  Day  3 f o r EXP2A  d i f f e r e n c e between t h e and  ft  in situ  surface  langley  f o u r t h week { F i g u r e  constant  Chlaiays  Environment  t o 550  a v e r a g e SB o f  the  on  Descriptive  appropriate,  during  7  25...;.  i n Tables  S o l a r r a d i a t i o n a t the  D a y s 6 and  both  {incoming}  respectively.  considerably  substations  physical, for  13  summarized  Physical  cages c o n t a i n i n g  Communities  the  i n Figures  and  The  appropriate  v a r i a b l e s are  grazing  6.  2. ,  dynamics  productivity  Day  (Figures tanks  with  was  depth  The values in the p l o t a r e f o r i n c i d e n t SB, which was 10% h i g h e r t h a n SB j u s t above t h e s u r f a c e o f the w a t e r , a s m e n t i o n e d i n C h a p t e r 2. T h i s was a l s o t r u e f o r t h e ether graphs of SB d u r i n g E x p e r i m e n t s 2 and 3. 1  33  (Tables  2  and  3)•  The d i f f e r e n c e i n t e m p e r a t u r e  surface  and bottom d e p t h s was c a .  5  period  of  time during  high  maximum i n s i t u examination  SH.,  ht s a m p l i n g  temperatures  o f the d i e l  °C  during  were c a . , 21.0  temperature  showed t h a t '15.0 °C t e m p e r a t u r e s a t s a m p l i n g ca.  5 °C .-at t h e s u r f a c e  mid  and  averaged  bottom 28.3  experiments  the  stations  ppt  and  by  was  late  ,  an days  increased  by  and 2-3 °G a t t h e  afternoon.  essentially  although  on two s u n n y  time  stations,  extended  The  constant  salinity  during  the  (C?=2.2%).  Nutrient Initial  and o u t f l o w  the  the experiments,  °.C  variation  between t h e  Conditions phosphate c o n c e n t r a t i o n s  relatively  constant  N ;P  inflowing  seawater i n d i c a t e d  nutrient  for  the  primary  averaged  atomic that  ratio  of  of  -  1  and  8.5  f o r the  the  limiting  particulate  organic  nitrogen  formation  2*; 15 uM P I  was  matter., , The with  greatest source  o f n i t r o g e n was i n the form o f n i t r a t e ,  inflow concentrations  27  ua N 1-* r a n g e  (CV=19S).  in  i n f l o w N.Q3 between t a n k s  18.5 uH N 1-* w i t h i n an 11-  T h e r e was no s i g n i f i c a n t d i f f e r e n c e except  However, t h e f l u c t u a t i o n s i n t h e i n f l o w  to;a  decreasing  tidal  inflow  first  reason.  and t h e g e n e r a l  the  the  levels  real  in  during  nitrate  were  increased  averaging  decreasing  trend  height a t sampling.time  to  week  when  Tank B f o r some NO3  with  time  c o u l d be a t t r i b u t e d * during the f i r s t  * The i n t a k e p i p e f o r t h e I n s t i t u t e d s e a w a t e r s y s t e m i s h i g h e r i n t h e water c o l u m n a t l o w t i d a l h e i g h t s and s i n c e N03 i n c r e a s e d with depth, t h e i n f l o w contained lower n i t r a t e c o n c e n t r a t i o n s .  34  part mass  of  the  e x p e r i m e n t and  (high temperatures  concentrations), Bay in  d u r i n g the situ  stations,  both  experiments  recovery  to  i and  both  due  week o f  to  the  Day  6,  fourth  with  levels  with  and  uM  The  N i ~  1  l e s s than  - 0.7  t o the s o u r c e  oysters  or  high  initial  were d e p l e t e d  at  N l - i during  1 UM  was  a  minor  c o n c e n t r a t i o n s a t the  were low  NH3  of  (max=1.85 uM  remained  scallops  was  uM  N 1~*  n i t r o g e n f o r the a t low  N lrr* ) i n d i c a t i n g  uM  )  There  an a v e r a g e c o n c e n t r a t i o n o f 0.74 little  nitrate  conditions i n English  17)..,  nitrate  low  water  end  of  N  1-*  weeks.,  production tanks,  •ca.a,- 0.4  (22.5  16  non-limiting  cf a surface  experiment.  averaging  (Figures  influx  unstable  ammonia c o n c e n t r a t i o n s  contributed  the  by  s e c o n d and Inflow  ),  oxygen  nitrate concentrations  all  the  and  probably  last  a l s o t o an  that  taken  steady any  up  (SD=0.42)  system.  In  state levels  excreted  immediately  NH3 by  ( by  the  phytoplankters.  Phytoplankton The including  Dynamics  dynamics the  productivity  of  the  phytoplankton (PBOD)  standardized  per  i n F i g u r e s 18  to  unit 25.,  and  resultant standing the  of standing  primary stock  primary stock  communities, (CHLA),  productivity (ASS),  are  primary rate  illustrated  35  Standing In  the  oyster  phytoplankton followed Day  Stock  (  tank  ca.  by a s u r f a c e  6  (Figure  (A),  46  ug  a  Chi  subsurface  j 1-* ) d e v e l o p e d  maximum o f o n l y h a l f  18) .  This  maximum  indicated  this  that  by Day 5,  concentration sinking  t h r o u g h t h e w a t e r column was a s i g n i f i c a n t  in  the s p a t i a l  and  a t t i m e s was g r e a t e r  During  the  values the  grazing  distribution  o f the primary  than the u p w e l l i n g  period  (t>6),  CHLA  o f 3.3 ug C h i a 1-* a t t h e s u r f a c e ,  mid  depth*  and  6.8  ug  steady-state  1-5 ug C h i a 1-* . the  with  station  significant  increase  (Day  I r * at  C h i a 1-* a t t h e bottom  station.  biweekly  19).  i n the inflow  6)  H03  Chi  during a  1-*  and was s i m i l a r  corresponding significant  the g r a z i n g 1.6  period increase  algal  of c a .  grazing  period  bloom o c c u r r e d  varied  at  i n EIP2A.  inflow  some  The  the  extent  standing  (t>10) a v e r a g e d the value  was p r i m a r i l y  i n CHLA t o l e v e l s of high  to  pressure.,  i n Tank B  This  at  was two d a y s l o n g e r i n  ug C h i a 1~* more t h a n  subsequent t o the period fourth  a  i n magnitude t o t h e o y s t e r  The d u r a t i o n o f t h e bloom  ,  assumed  concentration.  (8), the i n i t i a l  Tank B i n t h e a b s e n c e o f any stock  stock  periodic oscillations  t a n k . However,; t h e p o s t - b l o o m d y n a m i c s (Figure  to average  a t t h e end o f f o u r weeks i n r e s p o n s e t o t h e  In t h e s c a l l o p tank time  m/day.  !  A s e c o n d a r y c h l o r o p h y l l maximum o c c u r r e d  bottom  same  factor  4.2 ug C h i a  E x c e p t a t t h e bottom s t a t i o n , t h e p h y t o f l a n k t c n quasi  the  community  r a t e o f 0.25 decreased  on  of  phytoplankton determining  of  of ca.  5.7 ug  during the  due  to  13 ug C h i a  N03 a t t h e  end  of  the l  _  l  the  week, a n d , a s m e n t i o n e d l a t e r , t o t h e r e m o v a l o f f l o a t i n g clumps a t t h e s u r f a c e  of the tank.  36  Oxygen The  oxygen  production  of  oxygen  effective of  ca.  damping  17 mg  between  2),  averaging  and  of  averaging The scallop  l o w e r , and the  end  were  result  one-half  1~*  the  to  2Q  was  high  with  the  a  of the  the  production  the  initial  in  OXY  periods  initial  the bloom  occurred  of  thermal  p e r i o d , the  between d e p t h s  Cv=24J5.  The  phytoplankton  mean (Table  tank  experiment, reaching  OXY  a  similar  was  a maximum bloom  pattern  and  in  the  a t depth  were  ( F i g u r e 2 1).  At  concentrations  occurred  below  respiration  in situ  not evident  during  illustrates  experimental  exhibited  average  high  both  14058 a t a l l d e p t h s .  temporarily  due  to  net  with time  o f t h e f o u r t h week, o x y g e n l e v e l s a t t h e  of the  stress  mg  larger variances  reduced  probably  Figure  with  most of the  curves but  a positive  v a r i a n c e s were s i m i l a r  approximately  tank,  (CV=65$)  differences  f o r the t o t a l  during  oxygen  and  inflow  w i t h an a v e r a g e s a t u r a t i o n  1  tank,  Significant  during  21 OX  ca.„  the  o f the oxygen l e v e l s a f t e r  c a . , 11.6  supersaturated  oyster  in  constant 1~  coincident  although  concentrations  mg  maintained  1—* -.  depths,  stability,  7.5  Sfithin the  (OXYB) was  (OXX)  relatively  averaging  82%.,  of  concentration  s y s t e m s was  experiment, level  Levels  inflowing  mid  s c a l l o p s died during  station  concentrations,  r a t e s of t h e s c a l l o p s a s a  temperatures.,  at t h e  bottom  A  s t a t i o n , but  similar  oxygen  as n o t e d  later,  IXP2B a t t h i s  depth.  37  Primary P r o d u c t i v i t y The  primary  similar  pattern  ranged from during  productivity  to the standing  110 t o 170 ug C l ~  the i n i t i a l  A  similar  23),  as  a  concentrations.> between nitrate scallop  bloom  tanks  but  on  phytoplankton  from  A  followed  a  a n d t h e maximum  values  the surface  bottom  to  p r o d u c t i v i t y was e v i d e n t i n were  the  ca.  higher  difference  3535  1~*  previous  )  in  day;  in  the  this  greater,  standing PfiOD  c n Day 10 as a r e s u l t  (26 uH K  the  4  rates  of  stations  concentration tank  the  significant  and  -  o f primary  result  A  stocks,  hr  l  f o r Tank  (Figure 2 2 ) .  pattern  Tank B ( F i g u r e partially  curves  stock  occurred  of the high  inflow  to the  suggested  that the  p r o d u c t i v i t y was a l r e a d y n u t r i e n t - l i m i t e d by  this  time. The  standardized  considerably from  range  was C  of the o s c i l l a t i o n s  greater  (ug C h l a ) - *  intracellular productivity  During  h r  at this  time.  in situ  nitrate  nitrogen  levels  Day 8.  (Figure  4  ) and an  the  in  a situ  lower  influx  of  initial  not  The s i g n i f i c a n t  H03  ranging  p e r i o d , the i n the i n the  i n ASS between s t a t i o n s of c a .  bloom  9.0 -ug.  indicated  sere depleted  limiting  ,  primary  i n c r e a s e i n ASS on Day  15 a t a l l s t a t i o n s was u n p r e d i c t a b l e , s i n c e was  24),  increase  concentrations were  fluctuated  a decrease  The maximum v a l u e  -  on  -  (ASS)  the g r a z i n g  The v a r i a b i l i t y  h r * a t t h e end o f  although  EXP2A  was m i n i m a l , w i t h  (ug C h l a ) - *  {to two w e e k s ) .  also  that  (ug C h l a ) - * h r ~ * .  (2.5-7.5 ug C  period  productivity  a t a l l stations during  t-9 ug C  damping  primary  at t h i s time,  (plus a temporary  n i t r a t e l e v e l s ) and Sfi a n d TEMP were l e s s  there  i n c r e a s e i n the than  on  Day  38  12-  This  vitamin, week,  indicated  was  productivity at  solar  was i n Tank  the  A,  as  a  During  at  function  relatively higher  with  light  ASS  most  to  of  since  the  the rates  d e p t h would have a  of  SB,  w h i l e compared  productivity  rate  the  low  l e v e l of  to  mid  physically,  at  were a t  compared  to  the  a  primary and  incoming  mid  the  station;  bottom  station,  productivity  surface  function  a  fifth  period  higher the  as  surface  the  phytoplankton  as  the of  post-blccra  the  isolated  During  limitation  greatest  productivity  reasonable  phytoplankton  of  bottom,- a t t r i b u t a b l e  highest, standardized seems  productivity.  evidence  radiation.  this  p e r h a p s some s i c r c - n u t r i e n t , s u c h  l i m i t i n g primary  there  •least  that  which  would  cf  was  have  the  a  limiting  n u t r i e n t (s) . . The  standardized  unstable, undamped of on  as  ASS  confirmed  the  the  variances.  that  l e v e l of  during  depths;  the  costatum with  The  the  EXP2B was  rate  Figure  a large  i n the 25.  amplitude high  scallop  The and  value at  high inflow  nutrient  N03  5.6  ug  C  (ug  ..  ,  bloom (  en  in  ca.,  spp  was  exhibited  bottom  out  station  the.previous  day  average value  of  Chla)-*  hr~*  despite  for a l l the  high  • '  y  the  both 60S  other diatoms  N i t z s c h i a ••  curves  the  l i m i t a t i o n . , The  ca.  tank  considerably  a v e r a g e s between t a n k s were s i m i l a r  Composition of The  in  between d e p t h s .  10  reduced  illustrated  o s c i l l a t i o n s of  phase Day  productivity  .  by  Phytoplankton tanks  numbers)  ( Chaetoceros )  was  and  and  Comitucity  dominated  by  Thalassiosira  spp. .  ,  Navicula  nano-f l a g e l l a t e s  Skeletonema spp. , spp. .,  (20%) And  contributing  39  approximately Tank - A, During  Chaetoceros  the f i f t h  Navicula was  10% e a c h . .  sp.  third  sp.,,-  became  was removed f r o m recovery  sp.  the  feces, floated  worsened  and on Day 28, t h e c l u m p s o f N a v i c u l a  the an  surface  The  results  summarized three  meat  i n Table  weight  (JSETWH)  weight  calculation  weight  (PEE WM)  growth  total  weight  (NETWS) • , O n l y  the p r o p o r t i o n o f l i v e  greater  There  measurements  The p r o b l e m  19,  of  during  was  EXP2A EXP2A  (NET!T),  the percent  shell  are  i n the  meat  weight  increase  there  i n the  in  was no way  cultch.  The  p e r week was b a s e d on  the  linear  was a c o n s i d e r a b l e  significant  from  there  during  dimensions  growth  of the  range i n s i z e  t h e c u l t c h . , A few c u l t c h had o y s t e r s  was  contained  t h a n 2.0 cm., I t was i m p o s s i b l e t o  ".b-fej^i^i>-.-5 cm. , The NETHJH:NETfiS r a t i o •;  qiqas,  c o u l d be c a l c u l a t e d s i n c e  However, t h e r e eefl  -  the  stock.  o f NETWM and NETWS p e r o y s t e r  ^t iXa'e«!urate  •4«Sfe5n*l^''!'  oyster  During  4, which i n c l u d e s t h e n e t i n c r e a s e  number o f o y s t e r s  i  the  variables  and s h e l l  of estimating  the  of  Cragsostrea  there  were removed  i n Figure  of the phytoplankton  Growth o f the H e r b i v o r e s ,  B.,  o f t h e tank.  o f Tank B., As i l l u s t r a t e d  ifflmediate r e c o v e r y  However  which a l s o  scallop  benthic  stock.  i n Tank  clumps o f t h e a l g a e ,  to  phytoplankter.  pipe.  c f the phytoplankton  on t h e s u r f a c e  oysters  filamentous  the inflow  was a l s o a p r o b l e m  week, f l o c c u l e n t  the  the dominant  week, a t h i c k mat o f  ho s i g n i f i c a n t Navicula  After the addition of  with  within lengths  was a l s o c a l c u l a t e d . of  a l l the cultch  EXP2A.;  The p e r c e n t  i n c r e a s e i n meat w e i g h t  36.H6%4;ii  The h i g h e s t  average r a t e f o r t h e  ranged frcm  four  during  10.9% t o  substations  was  40  25% a t t h e mid d e p t h , a l t h o u g h t h i s significantly variability  greater  than the other  between SUBSTN.. The  p e r o y s t e r p e r week was value  for shell  g/zop/wk t o 1.17 NETWM:KETBS and  least  from  since  increase  (SD=-051 g ) •  0.83  g/zoc/wk> was  considered  i n meat  The which  ranged from  highest  t h i s r a t i o i s a f u n c t i o n o f the s i z e  Growth o f t h e H e r b i v o r e s ,  Chlamys  i s not  in  average  (SD=.G37) a t the bottom  of o y s t e r s per c u l t c h , a large variance  weight  corresponding  a l s o a . l a r g e range  t o 0.40., The  0.32  n e t be  d e p t h s i n view o f t h e  g  0.16  was  two  average  g/zoo/wk., T h e r e  variability  However,  0.24  w e i g h t was  ratio  mean c o u l d  0.5 the  ratio  station. and  number  unexpected.,  hastata  hericia.  during  EXP2B The scallop  results  of  population  than h a l f o f survived;  the  of  The  percent  increase  (NSURV)  and g r o w t h o f t h e  a r e summarized  scallops  at  the  i n Table  four  mid  5. , L e s s  substations  s c a l l o p s t h a t d i d s u r v i v e , most l o s t  e x c e p t i n Cage 2 w h i c h (HGTT).  survival  d u r i n g EXP2B 32  the  the  had  a  1.1%  increase  s m a l l e s t s c a l l o p s i n Cage in  length  (0.5%)  1 had and  comparison, a l l of t h e s c a l l o p s i n the f o u r  in  weight,  total  weight  the l a r g e s t average i n width  (2.0%).  cages at the  bottom  d e p t h s u r v i v e d . . However most s c a l l o p s d e c r e a s e d i n t o t a l and 10.5%  the  f o r Cage As  grazing the  average 12  mentioned period  mid  difference  depth.  percent  loss  r a n g e d f r o it 5-0%  (which c o n t a i n e d  However,  increased  to  ca,<  f o r Cage 9 t o  temperatures  °C a t t h e bottom during  weight  the l a r g e s t s c a l l o p s ) .  e a r l i e r , the average were 15.6  In  periods  5 °C .  The  during  d e p t h and of sclar  high  17.2 SR,  the °C a t the  radiation also  41  decreased s i g n i f i c a n t l y  with depth.,  SH or,TEMP, o r more, l i k e l y probably  responsible  Further  Investigation  T h e r e f o r e , high  a combination  values  of both v a r i a b l e s ,  f o r the low s u r v i v a l  and growth  o f t h e O y s t e r Food  Chain  of were  rates.  a t an I n c r e a s e d  Herbivore Density  EXP3A was i n i t i a t e d EXP,2A,  except that  24 , o y s t e r  and c o n d u c t e d  the i n s i t u  cultch,  using  8  substations.  with z o o p l a n k t o n a f t e r  was r e s t a r t e d  on A u g u s t  of the delay.  the  same  manner  as  h e r b i v o r e , d e n s i t y was d o u b l e d t o  contaminated  because  in  The  tank  became  a c o u p l e o f weeks and EXP3A  20 and c o n d u c t e d  for  only  three  weeks  The o y s t e r c u l t c h iwere added t o t h e t a n k  on Day 6 a t t h e a p p r o p r i a t e s u b s t a t i o n s , a s o u t l i n e d  in  Chapter  • .' |. j j • :  2.  D y n a m i c s o f t h e P r i m a r y Community The  flushing  the experiment and  primary  remained  (CV=1%).  c o n s t a n t a t 0.25 day-* d u r i n g  The r e s u l t s  variables  summarized i n T a b l e  the  rate  are i l l u s t r a t e d  Environment  Although  t h e w e a t h e r was b e t t e r  average  incident  daylength  temperature^  chemical  i n F i g u r e s 26 t o 32 and  6.  Physical  shorter  f o r the physical,  solar  (Figure  , during  radiation 26).,  was  However,  t h e r e was no s i g n i f i c a n t  EXP3A  than  EXP2A,  11% l e s s due t o t h e in  difference  terms  of  the  i n TEMP between  H2  experiments*; for the  The  reduction i n solar  r a d i a t i o n was  by a n i n c r e a s e i n t h e t e m p e r a t u r e o f t h e i n f l o w t o 16.2  °G  considerably  average., better  As  mixed  shewn  i n : Figure  during  EXP3A,  afternoon. ,  Nitrate the f i r s t  the  period. the from 12.9  were  were  even  during  c o n t i n u a l l y monitored  (SD=2. OS) which was  not  Inflow  The i n f l o w N03 was p r o b a b l y  the  during  averaged different  high  the l a s t  .inflow  week o f  temperatures  11 on, a n d t h e s u b s e g u e n t low i n f l o w c o n c e n t r a t i o n o f In-situ  t o v a l u e s l e s s than  although  nitrate  1 uM N 1  concentrations  (Figure  by t h e end o f t h e e x p e r i m e n t , h i g h i n s i t u  levels  recorded  after  a t a l l s t a t i o n s u n l i k e EXP2A.  Low i n f l o w  0.50 uH N l ~ * c o n t r i b u t e d l i t t l e  nitrogen  production  source  concentration  were  t h e blcom  _ 1  concentrations averaging  the  was  (SD=3.62) f o r t h e same  low d u r i n g  by, t h e  levels  .significantly  EXP2a a v e r a g e o f 21.8 uM N l-»  uM N l-» on Day 20,  depleted 28),  concentrations  e x p e r i m e n t , as evidenced Day  27, t h e t a n k  fieqime.  twelve days of the experiments  21.0 i uH N 1~*  maintain  •  Nutrient  from  compensated  f o r the  system.  The  a t any d e p t h was n o t s i g n i f i c a n t l y  i n f l o w v a l u e a t any t i m e d u r i n g  Phytoplankton  ammonia t o the  in situ  NH3  different  from  the e x p e r i m e n t .  Dynamics  The: d y n a m i c s o f t h e p h y t o p l a n k t o n  community  during  EXP3A  43  are  illustrated  primary  Figure  p r o d u c t i v i t y curves  approximately and  in  a  from  period  p e r i o d , CHIA was  with  the p h y t o p l a n k t o n  phytoplankton apparently  stock  also apparent  between d e p t h s was ,  During  averaged  grazing  i n the i n f l o s  i n t h e oxygen  o n l y 7.04  mg i  -  1  EXP2A, t h e maximum OXY  3.0 mg l - i  in  compared  the  less,  grazing  7,3 ug C h i a  pressure.  period  l-i in  However, t h e , which  was  source.  Levels s e a w a t e r c o n d i t i o n s was  concentration with  oxygen: l e v e l s f o l l o w e d a s i m i l a r  half  36% h i g h e r  was p r i m a r i l y C h a e t o c e r o s s p .  lack of constancy  averaged  stock  with  on Day 4,  r e j e c t e d by t h e o y s t e r s a s a s u i t a b l e f o o d  Oxygen The  oscillations  5.2 ug C h i a 1-* f o r t h e c o r r e s p o n d i n g  EXP2A, i n s p i t e o f t h e i n c r e a s e d  s t o c k and  The bloom o c c u r r e d  13,9 t o 16.2 ug C h i a 1~*  (t>6),  compared  the standing  e x h i b i t e d damped  EXP2A, and t h e d i f f e r e n c e i n GALA  ranging  as  Both  one week p e r i o d .  during the pre-grazing  with  29.  (Figure  a CV o f 10.2%.  30),  Although  p a t t e r n o f damped  was a p p r o x i m a t e l y  i n situ  oscillations  c o n c e n t r a t i o n o f 14.1 mg 1  l o w e r and t h e p e r i o d  which  a  _ 1  was c a .  week,  one-  t h e d u r a t i o n i n EXP2A., T h e r e was no s i g n i f i c a n t d i f f e r e n c e  oxygen l e v e l s between depths,  was  apparent  days  previously.  was 9.92  mg  production of EXP2A. ,  and a h i g h  between t h e p r o d u c t i o n  i~ l  The average v a l u e which  o f oxygen  positive  correlation  o f oxygen and CHLA o f OXY  represented  (OXYN) compared  a with  from  two  during the experiment 50%  decrease  tiefirst  in  three  the  weeks  44  Primary The  primary  throughout hr—  ,  1  Productivity  the  and  1.2  experiment.  experiment  31), averaging  also  s t o c k . ,•  exhibited during  were s i m i l a r between  were  phytoplankton (ASS)  productivity rates  positively  The  undamped  the  (Figure  oscillations  pre-grazing  During  period  to  the grazing:period,  the  the  high  vitamin  B12,  suggested  was  experiment, as well EXP3A . and occurred the  EXP2A  ugC  limiting  ASS s a s g r e a t e s t  was  period  period,  (ug C h l a )  - 1  hr-  1  Composition  was  grazing  system  in :  the  The  perhaps in  this  o f ASS between  initial  value  maximum  o f 2.4  during  During  the  p r o d u c t i v i t y r a t e averaged 25S l e s s  than  the  4.3 mean  i n EXP2A.,  o f jthe. P h y t o p l a n k t o n  Community  c o m p o s i t i o n o f t h e p h y t o p l a n k t o n community d u r i n g  similar  found  2., T h e p a t t e r n  , which was a l s o  EXP3A  t o EXP2A, w i t h S k e l e t o n e m a c o s t a t u m d o m i n a t i n g t h e  bloom a n d C h a e t o c e r o s s p . , the  nutrient,  was 2 5 % l e s s t h a n i n EXP2A.  time p e r i o d  at  a t t h e end o f  productivity  different.  the standardized  for the corresponding  The  very  from  was n o t l i g h t - l i m i t e d .  another  primary  as Experiment  productivity  8.8 a t t h e end o f t h e  on Day 3 i n EXP3A, and t h e a v e r a g e  pre-grazing  grazing  that  the  3 2 ) , ranging  i n s i t u n i t r a t e concentration  experiment a l s o  with  primary  (Figure  bottom s t a t i o n i n d i c a t i n g t h a t t h e system However,  27.2 ug. C 1"'  correlated  standardized  depths  period.  the  dominant  phytoplankter  during  L a r g e numbers o f n a n o - f l a g e l l a t e s  were n o t  e i t h e r EXP2A o r EXP3A i n c o n t r a s t (F£=0.25  day  - 4  )  with  no i n s i t u  to a similar herbivore  culture  population  45  (Brown a n d P a r s o n s , inflow  1972).  The b i c m a s s o f N a v i c u l a  pipe  was l e s s  the  oyster  cultch.  very  c l e a r a n d l a r g e amounts o f f e c a l  the  i n EXP3A a l t h o u g h By  The stocking The  g/zoo/wk EXP2A. .  growth density. maximum  m a t e r i a l were a p p a r e n t  at a Higher Stocking growth d u r i n g  d e n s i t y was d o u b l e d f r o m average  change  on  on  (SD=. 159) compared However,  the  during  of  EXP3A  average  any was  values  significantly The  same  greater problem  NETHMzNETSS r a t i o ,  with  large  The v a r i a b i l i t y  ratio  of  less  temperatures during promoted  an  the surface  0.24  g/zcc/wk  variability  due  was  0.16  (SD=.051)  for  within  substations  probability  that  t o the increased  the  stocking  between d e p t h s  and  a t t h e bottom  station  g/zoo/wk  g/zoo/wk  than the s u r f a c e with  from .03 t o This  value  during  growth i n s h e l l  NETWS  the  were  c r mid s u b s t a t i o n s . ,  variability  (SD=0.70)  o f 0.23  (SD=.3"76) f o r  the f o u r t h and f i f t h  increased  i n Table  was a l s o h i g h  which ranged  0.29  EXP3A, i n w h i c h t h e  EXP2A, a r e summarized  significant  (SD=.285) f o r t h e 24 c u l t c h .  average  Density  i n HETHJ3 per o y s t e r p e r week  (SD=. 178) f o r HETHH and 0.6O  .37  was some g r o w t h  t h e end o f t h e e x p e r i m e n t . Tank A was  r e s u l t s o f the oyster  removed t h e c h a n c e  not  on the  bottom of the tank. ,  Growth o f t h e O y s t e r s  7.  there  sp.  existed 1.00  f o r the  and  averaged  was h i g h e r EXP2A,  than the  The  week o f EXP2A may weight,  d e p t h w h i c h had a HETIHsBSXHS r a t i o  higher have  particularly at o f 0.26.  46  Farther I n v e s t i g a t i o n of the Scallop Flushing  Food C h a i n a t an  Increased  Bate o f t h e System  The  flushing  rate  and t h e ; t a n k s e e d e d  was s e t a t 0.75 day-*  a t 1500 h o u r s on J u l y  a s EXP2B.  As o u t l i n e d  i n Chapter  2,  introduced  i n t o Tank B on Day 11 f o r one  f o r Experiment  26, i n the same  the  scallop  3B  manner  cages  were  morth.  D y n a m i c s c f -the P r i m a r y Community During  the  constant  a t 0.75  seawater  system  through  the  experiment, day-*  the  (CV=9.4S)  and h i g h sediment  filter  to  the  at  sampling  time.  due  problems  to  with  the  loads reducing the flow  rate  The  flushing  b u t were r e a d j u s t e d  The r e s u l t s  F i g u r e s 33 t o 39 a n d s t a t i s t i c a l l y  Physical  r a t e d i d not remain  resevoir-  s o m e t i m e s d e c r e a s e d t o 0-6 day-* -*  flushing  rates  t o 0.75 day  o f EXP3B a r e i l l u s t r a t e d  summarized  in  i n T a b l e 8.  Environment r  Although to  the s o l a r r a d i a t i o n  decreasing  less  d a y l e n g t h , the v a r i a b l i l i t y  (CV=28%)  #  especially  at  was  15%  greater  EXP2B.J , B e c a u s e o f t h e in  situ  temperature  represented system  period for  increased  was l o w e r e d  net  i n SE d u r i n g EXP3B  (Days  was  1-6),  the  solar  EXP3B t h a n t h e same p e r i o d i n flushing  rate,  the  average  t o 14.8 °C a t a l l d e p t h s ,  a n e t t h e r m a l i n c r e a s e o f 3.4  ( F i g u r e 3 4 ) . . The  due  t h e beginning of t h e experiment  ( F i g u r e 3 3 ) . . D u r i n g t h e bloom radiation  was 8% l e s s t h a n i n EXP2B  increase,  °C f o r t h i s in  FXP2B  which  production  was  5.5  °C  indicating decrease  that  tripling  i n the  in situ  d i u r n a l i n c r e a s e was  Nutrient Inflow was  not  less  l e s s than  Day  6  and  recovery (Figure  concentration  was  situ  by  the  only  uptake o f  Standing The  time.  The  averaged  averaged  0.42  t h a n EXP2B, and  0.3  of  N  the  nitrate  average  N 1-*.  were tank  that  a  NH3  phytoplankton  no  again with  no the  ammonia  significant  steady-state and  was  during  inflow  T h e r e was  indicating  which  variance  concentrations  The  by t h e  uM  N I T * , i n the  uH  limiting  uM  19.3  existed  excretion  of  p e a k e d on  Day  However,  a  occurred  on  Dynamics  Stock  standing  concentration,  is  stock,  measured  illustrated  6 c o i n c i d e n t with larger  Day  secondary  stock  maintained  the  a quasi  the 36.  depletion  maximum of c a . an  During  displayed  as  in Figure  10 i n r e s p o n s e t o  concentration.  phytoplankton and  °C  the s c a l l o p s .  Phytoplankton  nitrate  2.0  /:  concentrations  35).  c h a n g e i n NH3,  'between NH3  higher  from  experiment  in  sampling  at  °C  concentrations  (CV=10%). < I n - s i t u  significant  2.5  caused a  fieqjme  nitrate  by  rate  flushing  temperature  significantly  depleted  the  the  initial  bloom  of  in situ  N03.  ug  hr  40 in  grazing of  level  a  The  increase  a series  steady-state  chlorophyll  damped at ca.  C the  1—*  - 1  inflow  period,  the  oscillations 30  ug C h i  a 1  48  -*,  and  there  The  dynamics of the  rate  of  0.75  was  day-*  EXP2B  (FB=0.25  ug C h i  a  three  times  1-*  effect system. and  i n fact,  total  experimental CHLA v a l u e  increased grazing  FB  in  pressure  f o r EXP3B  p a t t e r n from the  was  between  11;45  mg  t h e oxygen c o n c e n t r a t i o n initial  second, a s t e a d y - s t a t e duration during  of  EXP3B,  the  approached  grazing the  concentration and  50%  directly  During  the  that  the  dominated  a decrease  OXY  the  i n CHLA,  scallops  may  showed a  completely  during  There  have  EXP3B  average  EXP3E and  there  the  fclcom  the  period  a  standard  for  the  25%  the  inflow;  l e s s during  grazing  lew  period.  no  was  EXP2B:  70%  the  of  6.7%  deviation the  net  pre-grazing  less;  for  a v e r a g e CV  third, the  was  with  maintained  by  OXY  major d i f f e r e n c e s  compared  oxygen was  and  supported  The  were t h r e e  maximum d u r i n g of  i n EXP2B,  exemplified  value  during  results  depths.  l-»  during  level  (OXYN) was  greater  ( F i g u r e 37)  previous  d i f f e r e n c e between s t a t i o n s .  the  EXP2B and  indicating  the  23.7  approximately  rates.  cause  by  averaged  period,  5:1  during  productivity rate.,  the. reproducibility  first,  t h a n CHLA  experiment  d i d not MH3  higher f l u s h i n g  stock  during  depths.  levels  concentration  in  this  between  this  different  ratio increased to  oxygen c u r v e s  different  at  difference i n flushing  the  primary  Oxygen  stock  phytoplankton  t h e e x c r e t i o n of  the  The  The  average  to the  The  enhanced  by  day-* ) .  period,  difference  were c o n s i d e r a b l y  the  o f the  significant phytcplanktcn  f o r the  proportional grazing  no  which oxygen period  49  Primary P r o d u c t i v i t y The . 8 0  primary p r o d u c t i v i t y r a t e reached  ug C l-» h r  oscillations  to  -  on Day 6 and d e c r e a s e d  1  steady-state  rates  e x p e r i m e n t , PROD a v e r a g e d 38.3 ug C significant primary  less  productivity  during  rate  rate  bloom i n the  concentrations. low  of 1 . 1  and g r e a t e r  ug C  dominant  ;contrast  phytoplankter  during  Thalassiosira  sp.  More:significantly, Navicula  sp.  there  floating  were  no  standardized  period  due  days  to  the  hr-  1  during  value EXP2B  high  CHLA  constant  the grazing  at a  period  (6.0 ug C (Ug C h l a ) -  1  hr  (Figure 39).  Community  Skeletonema ccstatum remained the the experiment.  Other  diatoms  N j t z s c h i a sjap... .-. , N a v i c u l a  , and o n l y  was  was  during  EXP2B,  the  and i n c o n t r a s t t o EXP3B, ASS  average  were  there  The naximum three  of the Phytoplankton  numbers  and  1  o f ASS were r e l a t i v e l y  to  significant  -  During  the  (ug C h l a ) - *  variability  hr  38).  before  grazing  The v a l u e s  Ccmposition In  _ 1  occurred  E X P 2 B ,  i n c o n t r a s t t o the higher -*)  1  of ca  i n a s e r i e s o f damped  ^(Figure  d i f f e r e n c e between d e p t h s .  phytoplankton  a maximum v a l u e  a few f l a g e l l a t e s no  clumps  sp.- -  in and  were n o t i c e a b l e .  of  the  on t h e s u r f a c e o f t h e t a n k  filamentous a t any time  d u r i n g t h e EXP3B. ...  Growth o f the S c a l l o p s a t a H i g h e r F l u s h i n g The  r e s u l t s of the  population the  during  increased  survival  and  Rate  growth  of  EXP3B a r e summarized J i n T a b l e  f l u s h i n g i n t h i s experiment,  a l l of  the  scallop  9. ,, I n s p i t e o f the  scallops  50  died the  at mid  Only  the  surface  depth  the  and  substations, approximately  only  one  s c a l l o p s at the  growth d u r i n g was  initial  average weight of  i n Cage  12 i n c r e a s e d  the  s c a l l o p s at  increase  of  1.8%  by  only  the  g. 1.2J  bottom  i n length  with  the  shewed  percent  i n Cage 16.9  at  station  maximum  greatest  smallest  9,  bottom any  at  depth.  appreciable  , increase  averaging  died  in  16.8%  total  above  the  However, the l a r g e s t s c a l l o p s at  the  (Cage 8)  bottom  had  the  a corresponding  depth.,  largest  The  percent  increase of  0.5%  width. As  period in  bcttGm  EXP3B. , The  weight  in  Chlamys d i e d  one-half  pointed averaged  out  earlier,  14.5  °C  and  TEHP between d e p t h s .  light the dead  intensity  surface  .  within  resulting  frcm  was  the  week.. , the  environment f o r the  there  Therefore primary  Approximately a  the  was  no  the  significant  cause o f  the  of  the  10.0%  these  increased  m o r t a l i t y at  scallops  environmental  flushing rate  grazing  difference  i t appears t h a t the  one-half  Secondly,  increased  temperature during  provided  growth of C h l a m y s a t a d e p t h of  a  were  conditions favourable  1 metre.  51  CHAPSEfi  5.,  TWO-STAGE CONTINUOUS CUITUBES IN TUBBDLENT  8PWELLING SYSTEMS: DYNAMICS OF  THE  PBIMABY CCMMONITY AT  TWO  COMPABATIVE FLUSHING BATES The  dynamics  turbulent  systems  investigated flushing  of  natural with  phytoplanktcn  no  grazing  i n a s e t of experiments  EXP4A  and  1.00  During  the f i v e  relatively  at  0.49  the  experimental f a c i l i t i e s  in  Tank  angled  the  apparent  variable  measurements  stations  (surface,  After  each  taken  from  of  both  terminated  remove cn  i n t a k e a t the  Day  the i n f l o w  outflow)  was  the f l u s h i n g  any  and  experiments.,,  rates  remained, 1.00  day-i  interference  with  and  bad  this  set,  the  the other can  the  be  filtered  initial  samples  seeded  been c o l l e c t e d  The  tank..  seawater.  with  t a n k was  tube  in  tank  inflowing  through  to problems  pipe i n the  s t a t i o n and  in  rates  zooplankton.  35 due  Institute.  passed  during  - 1  differences  filled  t a n k s . . Then each  Bay  s e t of the  significant  tank  day  constant  the c e n t r a l sampling  i n c o m p l e t e l y mixed  and  of English to  0.5  <SD=.019)  12,  l i t r e s o f p h y t o p l a n k t o n s t o c k which  scree n  at  outside  toward  and  primary  i n f l o w i n g seawater  surface  r a t e s i n the  between the bottom  mid  e x p l a i n e d as sampling  were  flushing  period,  cn Day  became  Consequently,  '  The  day-*  (SD=.001). . However, a s a r e s u l t  first  two  were c o n t r o l l e d  week e x p e r i m e n t a l  were  compared  day-* d u r i n g EXPUB i n t h i s  constant  A  pressure  in  which  r a t e s o f t h e p r i m a r y system..  d u p l i c a t e primary systems  communities  with  10  from  the  a small diameter  wire  experiments  were  with the seawater  system  52  Dynamics  of  Flushing  the  Primacy  results  EXP4B) a r e g r a p h i c a l l y statistically  statistical and  time  illustrated  probability and  parameters  solar  langley  day  during  the  solar  of  a strong  (Table  solar  forcing  10).  radiation at  a  (Figure  station  otherwise  a  stated.  t o by t h e i r  summary  of  their  a  and  380 40}.,  day  - 1  T h e r e was no  coefficients  for  SB  p e r i o d i c i t y t o both primary systems A in situ tanks  temperatures (Figure  (TEMP)  41) and t h e  o f r ( A ) = . 7 4 8 and r ( B ) = . 804 between t h e  the  net  temperature  increase  s i g n i f i c a n t l e v e l for at least  temperature  in  Tank  (TEMPN)  one day i n b o t h  the lower f l u s h i n g  f o r TEMP and IEBEN w i t h  the  langley  from  l e v e l s and i n f a c t , t h e  correlation  The r e s u l t i n g  and  correlation  >0.99,  the  In the  factors  referred  ca.,  radiation  t a n k s , p a r t i c u l a r l y i n Tank A w i t h  was  15.  (SB) o s c i l l a t e d c o n s i d e r a b l y  averaging  serial  average c o r r e l a t i o n s  multiple  to  and  'significant' indicates  contains  s i g n i f i c a n t l y w i t h t i m e i n both  persisted  t o the  sometimes  experiments  the  high  (EXP4A and  .  ,  - 1  significance  varied  1  radiation  o f high  B  are  tanks  40 t o 56 10  l e v e l o f 0.05 u n l e s s  sustained period  and  Comparative  Environment  Incident  indicated  refer  and d e r i v a t i o n .  Physical  (SD=150)  Tables  and t h e term  names, and A p p e n d i x  description  i n Figures  in  »a' and. ••t*  respectively,  Variables  110r560  Two  f o r t h e two p r i m a r y  summarized  analyses,  statistical  computer  at  Bates  Experimental  data  Ccmaunities  station  rate.  The  and  time  A averaged  14.6 °C  53  (av SD=1.28)  #  temperature., temperature similarly of  1.0  to  13.0  the  a  net  A  pattern  of  a  correlation  and  «C  of  solar  significant  radiation  reduction  in  I t s h o u l d a l s o be  temperature  and  the  the  noted  salinity  of  ppt  variable part at  to the c o r r e l a t i o n  weeks than  of  .vicinity  experiments,  due  o f the seawater  correlated  in  °C  during  there  the  was  no  i n either  experiment  the  seawater  were  (av SD=0.40)  and of  ( T a b l e 10).  temperatures  Table  with the  10.,  N 1  to both  time  and t h e  was  varying t i d a l 4.  T h i s was  The  these  i n f l o w N03  solar radiation  However, a l t h o u g h  from  two  lower  d e t e c t e d by  during t h i s  three was  two  water mass i n  also  and o x y g e n l e v e l s  in  height  c o n c e n t r a t i o n was  of a surface  between  due  high during the f i r s t  the n i t r a t e  intake.  t a n k s were more  (C¥=14.5%),  — 1  i n Chapter  to the presence  the high c o r r e l a t i o n s  verified  the  reduced Although  temperatures  radiation  uH  N03  t i m e , a s mentioned  the  higher inflow and  between  height at sampling  average  .  slightly  inflowing  c o n c e n t r a t i o n s (N03)  w i t h t i m e , a v e r a g i n g 18.8  the t i d a l  °C  were  rate  fieqime•  nitrate  sampling  were  no s i g n i f i c a n t c o r r e l a t i o n  v a r i a b l e with i n c i d e n t s o l a r  Inflow  o f 2.6  a t 10.4  (SD=0.79) and t h e r e was  Nutrient  stations  higher f l u s h i n g  during  the  inflow  oscillations in  experiments,  that  the  amplitude  in situ  r e l a t i v e l y c o n s t a n t t o both t a n k s  either  and  decreased  of  from  between  c u t at t h i s  temperature  period  °C  periodic  (av SD=0.77), a n e t i n c r e a s e  nutrient-depleted  27.3  4.2  weekly  i n Tank B,  the average  1  average  tank;  high  apparent  day— ,  increase  days  the  period,  variables  also  the  are  significantly previously.  54  Although nitrate at  there  the  1.0  - 1  significant  flushing  day-i  the p o s t b l o c m level  no  between t a n k s , n u t r i e n t  t h e 0.5 d a y  at  was  depletion  r a t e i n Tank  flushing  period,  difference  rate  the i n f l o w  was a p p a r e n t  nitrate  presence  of  i n Tank B  (Figure  N03 r e m a i n e d  sporadic  i n Tank A a r e a t t r i b u t e d  concentration calculated The  of n i t r a t e  utilized  (NO3D), a n d i n c l u d e d contribution  of  uptake . o f t h e s e n u t r i e n t s  at  the  1~»  Stations  )  and  to  the  There  average  nitrate i n  concentrations  i n the s t a t i s t i c a l  ammonia and u r e a  i n f l o w c o n c e n t r a t i o n o f ammonia U  high  by t h e p r i m a r y  in either  of  mixing.  The  community  was  summaries.  as n i t r o g e n s o u r c e s was  no  significant  tank .due t o t h e low  average  (0.4 4 ufl N 1~* ) and u r e a  large  6  4 2 ) . „. D u r i n g  to the incomplete  f o r t h e p r i m a r y s y s t e m s was m i n i m a l .  by b a y  A w i t h a two day t i m e l a g  o f c a . , 19 uM N 1-* , b u t t h e r e was no i n s i t u  Tank B a n d t h e  uH  i n the inflow  variances within  (0.86  and between  during t h e experiment.  P h y t o p l a n k t o n - Dynamics  Standing The  phytoplankton  chlorophyll nutrient  a.  -  1  including  bottom CHLA  was p o o r  #  the i n i t i a l  on Day 6 ( F i g u r e  between  the  bottom  with  ug  Chi  18.3 ug C h i a  a  1~  t h e r e was a s i g n i f i c a n t  at a l l four stations  as  43) and f o r t h e d u r a t i o n o f  t h e h i g h e r v a l u e o f 20.5 Although  day  bloom was c o i n c i d e n t  (t=29), t h e s t a n d i n g s t o c k averaged  station.  with time  s t a n d i n g s t o c k was e s t i m a t e d e a c h  I n Tank A  depletion  the e x p e r i m e n t l  Stock  (av C V = 3 2 » ) ,  station  and  the  1  at  the  variance i n  the c o r r e l a t i o n other  ijj  situ  55  stations  for  CHLA, a s w e l l a s TEMP, N03 stock  the  other  fairly  regularly  on Days  17, 24 and 31 and t h e minima cn Days 21 and 28. „  major  differences  a one week p e r i o d ,  in  produced  subsequent  8,  to  due  this at  time.  the  Day  during  9  to at  with  the  depth.,  2.0  standing first  the i n i t i a l  total  times  Day  although  (t^27)  (P>.01)  was  different  15  except standing  the  the  standing  averaged  but s m a l l  40.6 ug increase  p e r i o d , the average 32.6  ug C h i a 1~»  ,  i n Tank A (0.5  t h e dynamics of the  were c o n s i d e r a b l y  Day  maintaining  Secondly,  FB)  was  on  higher  the c h l o r o p h y l l a c o n c e n t r a t i o n  Thirdly,  stock  - 1  bloom  nitrate  until  experimental  (1.0 d a y  several  phytoplankton  between t a n k s  in  the  few weeks, t h e t i m i n g o f t h e naxima and minima d u r i n g t h e  last  two  Tank  B  initial  weeks were s i m i l a r . exhibited blocm  oscillations the  time. ,  a significant  s t o c k i n Tank B  FB).  - 1  For  exhibited  12 due t o a p r o b l e m  this  oscillated  r a d i a t i o n and n i t r a t e a t  the n i t r a t e - d e p l e t e d p e r i o d was  day  Day  12. ...  maxima o c c u r r i n g  which had a s i g n i f i c a n t l y  C h i a 1-* and t h e r e  exactly  First,  The CHLA maximum d i d n o t o c c u r  from  standing  B.  bloom  low i n f l u x e s o f s o l a r  a r t i f i c i a l turbulence stock  Tank  sith  to the d e p l e t i o n of i n s i t u  t h e bottom s t a t i o n ,  stock  stations  Day  the s t a n d i n g  The d y n a m i c s o f t h e p h y t o p l a n k t o n  j  after  However  with  at  and OXY,  pattern  compared in  similarities  although  a  Tank in  the s t a n d i n g  However,  with  A.  This  stock  in  o f damped o s c i l l a t i o n s a f t e r  the  the  the  more  standing  unstable  was e x e m p l i f i e d  the  variances  between  stock  was t w i c e  as l a r c e  pattern  of  statistically tanks  for  i n Tank  B.,  by  CHLA  56  Pigment The (CHLC)  Batios  concentrations and  carotenoids  summarized i n T a b l e pigments a ratio  i n the (BA)  in  the  well  as  ratio  illustrated  average  1.12  between t h e  i n the  initial  pigment  and  STATIOH  C:CT  r a t i o s which  chlorophyll  was  c  The  pigment  a ratio  in Figure  44  ratio  from  t a n k s showed  during  time,  reflects  measurements  (see  45.,  for  this  0.105  The  but  in  The the  the  not period  the  seed  B respectively, a v e r a g e CTA  multiple  as  ratios the  1.12  correlation  i n d e p e n d e n t f a c t o r s TIME  f o r both tanks, except f o r the  probably  the  Figure  l i t t l e ' change from  population..  >0.90  the  and  ratios  ratio..  both  r a t i o s of  (t=29) , a l l p i g m e n t s r a t i o s  0.042 i n Tanks A and  r a t i o s and  are  (CTA)  different sith  CA  ratios  chlorophyll b:chlorophyll  pigment BA  (CBLE), c h l o r o p h y l l c  i n c l u d i n g the  period  significantly  decreases i n the  f o r the  the  12,  population.  t o 0.086 and  and  and  Table  a d e c r e a s e i n the  large  1..28  and  nitrate-depleted  population  of  also  stations.,  indicated  (CT)  carotenoid:chlorophyll  both tanks sere  between  11  seed  and  experiments are During  of c h l o r o p h y l l b  poor  precision  S t r i c k l a n d and  C:A  and  of  the  Parsons,  1972,  p.187).  Oxygen The as  a  Levels  oxygen c o n c e n t r a t i o n  state-determined inflow  p e r i o d . / I n Tank A,  the  oxygen  , a f o r c i n g v a r i a b l e as  variable,  (av SD=.675) i n t h e  in  (OXY)  OXY  (OXYK) were n o t  to  averaged  both t a n k s during  maximum and coincident  the  7.56 the  mg  l  -  *  experimental  maximum n e t  w i t h the  well  initial  increase depletion  57  of n i t r a t e on  Day  and bloom o f p h y t o p l a n k t o n  19  (  OXY {19) = 13. 35  ;a=3)  as a r e s u l t  solar  radiation  station, (t=29) increase  Tank  from  11.18  Days  18  mg  l *-  r  to  with  there  between s t a t i o n s , mean, f o r t h e  20-  during  remained  and  coefficient  h i g h a t >0.94  mid  the i n i t i a l  1-*  ;a=4), a l t h o u g h  1-*  occurred  net;  (Table  during  pattern  and  OXYN  was  grand  negative  The  multiple  with  d i d occur  both  i n Tank, B  on Day 8  factors  subsequent  ( OXYN(12)=5.65 mg  two l a r g e r maxima o f 5.91 mg l  (t>6),  Oxygen s a t u r a t i o n  the post-bloom  net  However,  -  1  and 6.35 mg  (Figure  46).  i n o x y g e n i n Tank B was 20'.% l a r g e r t h a n  was s i m i l a r  1  small  stations-  and  depletion of n i t r a t e  from about 79% t o s u p e r s a t u r a t e d 130%  -  14). ,  f o r t h e comparable time p e r i o d (av SD=0. 193).  a 3.71 mg l  d i f f e r e n c e i n OXY and OXYN  on Days 26 and 34 r e s p e c t i v e l y  increase  period  i n OXY and OXYN  stations  f o r OXY  A l a r g e i n c r e a s e i n oxygen to  post-bloom  s m a l l p o s i t i v e d e v i a t i o n s from t h e  surface  and  t h e bottom  t i m e b u t n o t between s t a t i o n s -  d e v i a t i o n s f o r t h e b o t t o m and o u t f l o w correlation  the  The v a r i a n c e  was a s i g n i f i c a n t  with  Excluding  which r e p r e s e n t s  i n o x y g e n i n Tank A.  B,  occurred  of the high i n f l o w n i t r a t e c o n c e n t r a t i o n s  highly significant in  4 6 ) , but  mg 1~* ,a=3; GXXN (19) =6. 47 mg 1-*  t h e a v e r a g e oxygen l e v e l was  (figure  averaging  i n Tank A  4.44  mg  (SAT) i n c r e a s e d i n b o t h  levels  averaging  The  greater  1-  1  tanks than  p e r i o d and t h e a n a l y s i s o f v a r i a n c e  t o t h e OXY a n d Q X Y N  r e s u l t s f o r both  tanks.  58  Primary Primary  Productivity productivity,  components,  was  measured  estimates  of  net  exudation  of  organic  measurements the  units  respiratory  using  particulate carbon  net  using  (R. Q.)  and  B,  averaging  respectively, ten was  TIMES  ug  particularly  in  occur u n t i l  for  PGO  with  in  ug  C  l  A.  * hr  during  - 1  *net p r o d u c t i v i t y ' ,  respectively. carbon-14  86%  of  However* method  o f 1.2  and a  high i n both  Tanks  6).  Although  stations, rates  multiple  relatively  due t o t h e three  for  there  there  with  the  was  time,  maximum d i d correlation  p o o r i n Tank A deviant  bottom  weeks t h e r e  was  a  ,  averaged  based  1  the  40  ug C 1  _ 1  hr-  1  and  EXP4A and EXP4B, and t h e p a t t e r n s  average  the  was  p a r t i c u l a r l y high r a t e s  and  and  and 340 ug C 1~* h r -  1  The  the f i r s t  (RES)  with  ! 82%  were  between  47).  tanks.  similar  was  ((P. Q.)  productivity  (r=.96)  ,After  repiration rate -  guotient  (PGC)  STATION  w i t h Tank B  Tank  (RES)  (PNG) , which were c o n v e r t e d t o  i n PGO  (Figure and  and  t h e o x y g e n method f o r  day s t a r t i n g on Day  h i g h e r c o r r e l a t i o n between The  (PfiOD)  (PGO), r e s p i r a t i o n  C 1-* h r -  i n these  15  TIME  (r=.78) compared station  fixation  Tank ., B i n w h i c h t h e p h y t c p l a n k t o n  Day  the  technique f o r  mid and b o t t o m s t a t i o n s  no s i g n i f i c a n t d i f f e r e n c e  a s i g n i f i c a n t variance  not  rates  220  third  of  o f 1.0. ,  f o r the surface,  (every  analysis  (IXC), plus  a photosynthetic  The.gross p r o d u c t i v i t y A  carbon  productivity  quotient  an  the r a d i o c a r b o n  of g r o s s p r o d u c t i v i t y  resultant  carbon  including  on Day  on t h e  12  oxygen  gross p r o d u c t i v i t y  the net p r o d u c t i v i t y (PBOD) , was  only  (Figure  were  48).  method  47  The  (PNO) ,  i n T a n k s A and B  rate determined  by  a sisall f r a c t i o n of  PGO  59  (0.21  i n Tank  large  a v e r a g e d i s c r e p a n c y between t h e two methods o f  •net  A and 0.17 i n Tank B) , i n d i c a t i n g t h a t  productivity*.  (McAllister e t a l . to  represent  as  constant a  also  suggested  ,1964; E p p l e y  and S l o a n ,  of cellular  during  both experiments difference  Daily  incubation  The a  rationale  function  estimates  lower l i g h t  late  afternoon  particulate  o f SR  factor  The  0.35 mg C I  was m u l t i p l i e d by the  day  t o the  (see A p p e n d i x 1 ) . since  maximum  PROD i s  -  1  SR,  daily  t o te proportional early  morning  to and  f i x a t i o n of  d y - * a n d 0.46 mg C  50).., I n t e r e s  the experiments,  of  2. J9 mg C 1 with  4.21 mg C 1~* dy~* on Day 12 i n Tank Values f o r the p r o d u c t i v i t y  the  l ~  l  were no major c h a n g e s i n t h e **  PGODY, v a l u e s a v e r a g e d  dy-* d u r i n g  (Figure  during  r e s u l t i n g estimated  productivity,  of  of  be e x p e c t e d  uith  1  was  8) and t h e p r o d u c t i v i t y  period  C productivity  -  days  was t h a t  l e v e l s during t h e remaining  time  was r e l a t i v e l y  (PCDY) t o n o r m a l i z e  period  (Chapter  the  could  carbon averaged  (EXC) a s  i n Tank A due t o t h e  during  the i n c u b a t i o n  periods.  into  49) a l t h o u g h t h e r e  of productivity  i n T a n k s A and B, and t h e r e  mg C l  rate  between  solar radiation  were measured d u r i n g  productivity the  radiation  The r a t e  during  appears  station.  for t h i s conversion  hyperbolic  - 1  solar  period.,,  radiation  rates  dy  in  r a t i o of the t o t a l  solar  (Figure  studies  carbon, t a k i n g  e s t i m a t e s o f P80D were c a l c u l a t e d  differences  other  due t c e x u d a t i o n  between s t a t i o n s  h i g h e r v a l u e s a t the bottom  the  carbon  was a  estimating  1965), PROD  r e s p i r a t i o n . ., The »*C p r o d u c t i v i t y  significant  any  by  the net f i x a t i o n of p a r t i c u l a t e  account the losses well  As  there  _  1  daily dy  a aaximum  - 1  gross  and 3.17  v a l u e (a=3)  B (Figure 5 1 ) .  component  variables  were  also  60  standardized  on a s t a n d i n g  represented  by  standardized  PGOST,  gross  although  particularly  i n Tank A  Table  higher  14,  basis  PNGST,  ( ug C  BESST,  p r o d u c t i v i t y (PGOST)  experiments,  and  stock  the  there  (ug C h l a )  ASS  and  ( F i g u r e 52).. average  r  and  i n i n Tank A and t h e r e  was  EXCST.  oscillated  was a d e c r e a s i n g  with  in  -  1  )  The  during  trend  As i n d i c a t e d maximum  hr  - 1  the time,  Table  13  v a l u e s o f PGOST were  more v a r i a b i l i t y compared  with  Tank B. All  t h e component v a r i a b l e s o f PGOST were a l s o  magnitude for  in  Tank A, w i t h  for  Tank B.  ug C  (ug C h l a )  each  system reached  initial  - 1  hr  -  standardized both  measurement decreased  The h i g h e s t  ) occurred  1  i t s maximum  bloom p e r i o d  between  net  primary  STATION times  in  54).  •  .  the  During  exudation  the  bottom  respiration  rate during  maximum v a l u e s  EXP4B, t h e h i g h e s t  2.0  BESS1-1.5, ASS=1.7  tacks three  BESST  values  time  and  (ASS=7.2  days  before  s t o c k l e v e l s d u r i n g the t h e post-bloom  productivity  with  standardized  with  standing  tanks  lower a t  BES,  o f 2.4 f o r BESST,  and T I H E . „ A l t h o u g h  for  both  in  assimilation rates  i n both  (Figure 53).  significantly  to  values  ASS and 6.5 f o r EXCST, compared w i t h  EXCST=6.0  the  average  greater  remained  there  rate,  station  constant  were o n l y  EXCST  and  the in  five  generally  values  Tank  EXE4A and EXP4B  o c c u r r i n g on Day  period,  A.  wa  s  were The  similar  12.,, However, i n  were a p p a r e n t c n Day  6  (Figure  61  Estimation Since  the  productivity on,  t=5,  -of P a r a m e t e r s t o r a P r i m a r y P r o d u c t i v i t y exudation  time,  a-3  to  rate  was o n l y  measured e v e r y  a productivity  component  evaluate  parameters  the  analysis in  Model second  was  the  based  following  p r o d u c t i v i t y sub-^model:  Gross  Productivity or  = Assimilation  1 = ASS/PGOST * RESST/EGOST + EXCST/PGOST  The r e s u l t s a r e shown i n T a b l e seems  a  good  estimated tanks  there  about  variables in  (ESTPGO)  flushing  rates,  60%  increased  components  t=5 during  particularly  are  were  with  of  N03/CHLA r a t i o  in  Tank  A,  inflow  gross was  ALPHAC  (  in  both  i n ESTPGO  spite  assimilation  calculated that  with  of  of  the  representing ca-  20%  to  The  assimilation  weeks of t h e  suggesting  that  concentrations.  productivity  f o r t=10 and a=3, t h e  these parametric  and f i f t h  (EEGQ)  was  fraction  experiments,  APGO also  estimates  increased the  lowest  in  exudation when t h e  highest.  P a r a m e t r i c e s t i m a t e s were a l s o slope,  t o 1.00  The  c f t h e p a r a m e t e r s f o r the  and l o s s e s  reasonable..  the fourth  response t o high fraction  model  t o e x u d a t i o n . , When t h e r e s p i r a t i o n and  v a l u e s were s i m i l a r , i n d i c a t i n g . on  additive  a r e s i m i l a r between t a n k s i n  r e s p i r a t i o n and c a .  based  i s close  the estimate  20% o f t h e g r o s s p r o d u c t i v i t y  assimilation  this  was no s i g n i f i c a n t d i f f e r e n c e s  TIME o r S T A T I O N , / S e c o n d ,  difference  15., F i r s t ,  a p p r o x i m a t i o n ., o f t h e e x p e r i m e n t a l r e s u l t s .  gross productivity  and  component  .Respiration. * Exudation  ug  C  (ug  calculated  Chla)~»  for  the  hr~* (ly/min)-*  initial ) i n the  62  hyperbolic the  productivity  measurements  questionable radiation  accuracy  at  was e s t i m a t e d in  of  depth  versus l i g h t  (PAB) r e l a t i o n s h i p .  the e x t i n c t i o n c o e f f i c i e n t due  to  faulty  (PABZC) used  using B i l e y * s  (EXTK)  equipment;  were o f  the  i n the c a l c u l a t i o n  f o r m u l a t i o n o f EXTK  Since  solar  o f ALPHAC,  (See  Chapter  2)  the following equation; PAEZ  The  (langley  average  (ly/min) Maximum  value ,  - 1  mn i r* )  was  ALPHAC  predictably  N03(IN)  above a v e r a g e .  stock  per  Tank  between  tanks  (ALPHAG)  1  significant  time  hr  net  significantly obtained  nitrate  since The  -  1  i n both  for  tanks  below  -  1  ALPHAG.  on Day 30  average  t h e s t a n d i n g s t o c k and  was  hr  and  nutrient  i n Tank A, t h e s t a n d i n g  equal. ,  Therefore  i t seems  both  predominantly  systems  had  slope  based  on g r o s s  f o r both  tanks,  averaging  1  , although with  time.  since A  i n both  depth  The s i g n i f i c a n t  surprising  f o r ALPHAC.  than  - 1  d i f f e r e n c e s i n ALPHAC  (ly/min)-  with  (ug C h l a )  were no s i g n i f i c a n t  initial  56).  C  lower  the l e v e l s  i n c r e a s e i n ALPHAG  (Figure was  Although  was v e r y s i m i l a r  (ug C h l a ) -  PABZO  of  that there  communities..  much  ug  SB a n d TEMP were  B were d o u b l e  unit  reasonable  11.0  v a l u e s o f 30.0 o c c u r r e d  55) , a t w h i c h t i m e  in  (PABZC/240.), * EXP (-EXTK*Z)  f o r ALPHAC,  (Figure  flux  =  similar  productivity  cases  64.3  ug  C  there  was a  due t o t h e d e c r e a s e i n  variability  both  diatom  ASS  and  analysis  i n ALPHAG PABZ  with  varied  of variance  was  63  Composition The  size  monitored The  composition o f  during  numbers  both  of  concentrations Parsons  o f the Phytoplankton  as  particles  (microns  the  phytcplarktcn  experiments  converted  m l ) , as  periods  similarity time..  unimodal  during  throughout variation  EXP4A.  The  first  the  the in  composition  trend  the  (Figures  57  and  experiment  (Figures  22  to  magnitude similar  over  time.  t o 61 a r e  i s the  curves  over  volume a t 18.4 58)  25)  persisted with  Secondly,  between s t a t i o n s ,  post^  note  size  maximum  size,  and  to  bloom  the was  with  57  bloom  point  i n t h e s h a p e o f t h e volume v e r s u s  The  microns  of  i n S h e l d o n and  Figures  r e p r e s e n t a t i v e o f t h e r e s u l t s d u r i n g both the bloom,  volume  of the p a r t i c l e  (microns)..  little  the  size  and t h e c o r r e l a t i o n  between t h e C o u l t e r c o u n t s and c h l o r o p h y l l a was g e n e r a l l y e x c e p t f o r a time l a g i n the p a r t i c l e  was  Counter.  into  outlined  as a f u n c t i o n  diameter  community  using the C o u l t e r  were  per  ( 1 9 6 7 ) , and p l o t t e d  measured  the  Community  good  volumes d u r i n g the i n i t i a l  bloom. The respects, the  results  f o r EXP4B  were  e x c e p t f o r two f e a t u r e s .  initial  bloom  were  higher  similar  The p a r t i c l e on  secondly,  t h e magnitude o f t h e maximum  was  t w i c e as l a r g e  about  Throughout  species  present  decipens  (?) , T h a i a s s i o s i r a  A  filamentous  12  i n most  volumes  during  than  volume p e r  t h e dominant  i n both systems.  were  Day  EXP4A  ml  Day 9, and in  EXP4B  as EXP4A.  the experiments,  Skeletonema costatum  to  mostly  b e n t h i c diatom  I n EXP4E, t h e  diatoms,  aestivalis  p h y t o p l a n k t e r was  including (?) and  other  few  Chaetoceros  Nitzschia  spp.  ( N a v i c u l a s|.„ ) was a p p a r e n t on  64  the  s i d e s of t h e p r i m a r y  was  low  the  compared  previous  with  production  the phytoplankton  experiments  benthic  non-turbulent  systems.  In  EXP4A,  diversity  flagellate species period,  were  the  post-bloom  , a Prorocentrum  s p e c i e s and  These f i n d i n g s are c o n s i s t e n t with  previous  laboratory  results  fluxes  tend  increase dominance  such  the  on t h e s i d e s o f  field  and  during  and a few  Navicula  Tank  T h e r e was  .sc.,  phytoplankters  also less  Tank ft t h a n  most  of  noticable  i n c l u d i n g Gymnodinium  some c r y p t o m o n a d s .  comparison, of  there  greater  In  production  in  a  stock..  stock  indicated that  d i a t o m s was s i g n i f i c a n t was  tank, hut the s t a n d i n g  B.  which i n d i c a t e d t h a t s y s t e m s with to  have  more  phytoplankton  i n the nutrient concentration of  diatoms.  of the phytoplankton the primary  community  a s the N a v i c u l a s p .  at  high  community  diversity,  r a t e s o f 2.0  washed  became d o m i n a t e d (Brown  lower n u t r i e n t and  an  c a u s e s an i n c r e a s e i n t h e  flushing was  and  by  and P a r s o n s ,  day  - 1  ,  out o f the system benthic 1972).,  diatoms  65  CH APTER 6. ,  THO-STAGE CONTINUOUS CULTURES  OF  PLANKTONIC  HE8VIVO ROUS POOD CHAINS: DYNAMICS OP T EE PRIMARY COMMUNITY VARIABLE FLUSHING A  set  of  investigated dynamics  two-stage  the  but v a r i a b l e ,  production  Tank  rates  respectively. due  to  were  altered  t o 0.50  different  to  On  rate  froa  experimental  experimental  either  continuous cultures presented i n Chapter During primary  the  function  weeks  continued rate of  stage o f the  A or B  was  This provided I  investigation  i s summarized  fed  i n Table  to  IV)  of  the two-stage  16.  and  four  The r e s u l t s  the  are  7.  were light  of t h e t e m p e r a t u r e  phytoplankton,  the  experimental  versus  day-*  t h e f o u r h e r b i v o r e t a n k s c o n t a i n i n g the  for  productivity  productivity  the  herbivorous  E was  Tack  c o n d i t i o n s (Experiments  design  6  the i n i t i a l  d e n s i t i e s o f o y s t e r s p l u s cne c o n t r o l .  unique  28  0.10  by two  day-* on Day 4 1 . , I n t h e s e c o n d  rates into  In  ( D i l e p t u s sp?) and t h e o t h e r  reset  t h e outflow from  and  and  i n p r o d u c t i o n Tack rate  using  (FR) was s e t a t  ended a f t e r  system  the  system. ,  14  day-*  i n t h i s tank  ciliate  the  were  case,  examined  Days  0.50  o f the primary  The e x p e r i m e n t  of  flushing  duration..  one a h o l o t r i c h  p r o d u c t i o n system,  three  experiments  were  rates  initial  Day 49, w i t h t h e f l u s h i n g  1.00- day-*  at  the  The e x p e r i m e n t  a hypotrich. to  A,  contamination  protozoans,  culture  communities  flushing  day-* f o r two weeks  flushing  continuous  primary  controlled,  0.25  BATES  i n t h e two p r o d u c t i o n s y s t e m s . , In t h i s  of  AT  period,  two  related  investigated. experiments and  the  were  nutrient  to e s t i m a t e the e f f e c t s o f these  A  aspects of series  conducted status three  of as  of  a  the  variables  66  on t h e p r i m a r y p r o d u c t i v i t y results  are  was  analyzed i n  also  verify The  presented  that  some  procedure  with  a  that  either Guillard^s  or  vitamin  the  same c o n c e n t r a t i o n  17)..  one  increase control..  at  level. the  h i g h light  same c o n c e n t r a t i o n  of  was  on  no  experiments  system.  radiocarbon  uptake  Tank  B were  aiix  of t h e  and  indicated  (Table  of v a r y i n g  light  the  other  that  only  the  a B12  to  the  i n c r e a s e i n primary which c o n t a i n e d  Therefore  limiting  at  significant  i n comparison  B12.  B12)  medium  with the F enrichment  the primary  no  were made a t  produced a  significant  vitamin  enriched  (with  F  two. days  rate  to  the  enrichments  intensity  and  productivity  limiting  from  level  productivity  day  H03  not  two  The r e s u l t s  However, t h e r e was  that  primary  other  the samples  conducted  on e i t h e r  probable  the  as t h e a d d i t i o n  were  procedure  enrichment  The l a t t e r  i n the primary  productivity the  to  The  was  at a l i g h t - l i E i t i n g  light-saturating enrichment  of  The  F medium, o r a v i t a m i n  only*  Experiments  intensity,  series  similar  except  B12  8.  i n Chapter  micro-nutrient  was  measurements  of the system.  nutrient  is  seems  during  these  at Variable Flushing  Bates  experiments.,  Dynamics  of  the P r i m a r y Communities  during the Experimental period  A f t e r each inflowing  primary  seawater  and  tank  *as  the f l u s h i n g  from  e a c h t a n k were t a k e n . , I n t h i s  tank  was  so  that  seeded both  with  filled  10 l i t r e s  experiments  rates of  the  set, i n i t i a l  filtered samples  experiments,  each  of phytoplankton stock from  EXP4B  would  set  with  have s i m i l a r  initial  primary  67  communities, other  composed  diatoms  and  (EXP5A*EXP5B) statistical  0.50  Physical  in  , and  0.10  the  experimental  in  period  i n Tank B,  langley  day-*  and; v a r i a n c e s  1,  of  langley  dayr*  2 and  six  the  incident  (SD=106).  very  during  second  p e r i o d and  radiation  variability  in  three  coincidence  alteration  i n flow  increased  TEMP) and  * EXP4B was  the  experiments  to  90,  In the t o the  with  following 0.25  SB  was  the of a  1.0, d a y  and  day  - 1  the  rate  averaged  400  i n F i g u r e 62, between  in  the the  340  l a n g l e y day-*  The  ccmhinaticn  flushing  (TEHFN) o f  a  three  langley  380  (with  means  EXP5A, a v e r a g i n g  1,  reinforcing  total  flushing  500  day-* (SD=49) of  the  rate r e s u l t e d in  6.7  °C  e x p e r i m e n t i n Tank A  day-*  lower than  two  - 1  radiation  rate, particularly  t o 0.50  terminated  includes  , 3.8  two-week p e r i o d s i n EXP5A.  i n TEHP d u r i n g  to the  was  18).  temperature i n c r e a s e s  8.8°C d u r i n g  the  Period  solar  in  ahich  rates  variability  t h e FB  few  rates respectively. ,  different  (Table  the  23.,  plus a  the  62  3 refer  illustrated  period  due  to  solar  flushing  (SD=15) the  As  were  variable  for  weeks,  during the t h i r d  was  18  f o r Tank A and  f o r SR  periods  and  Figures  day-* f l u s h i n g  first  period  average net  R e s u l t s of  Tables  EXP5A, P e r i o d * s  costatum  Environment  During  (SD=132)  *  illustrated  summaries  day—*  of Skeletccema  flagellates  are  d i s c u s s i o n of ,  mainly  effect during  °C  The  large  (Figure of  Period  subsequent  SB 2,  ,  63) with when  decrease  average., Temperatures reached  days b e f o r e  t h e . s t a r t . o f Experiment  a  5  68  maximum o f -2-1.9 ° C d u r i n g , compared  with  the  day-*  0.50  18.7  although  stations  was  significant (Table  20).  °G  The  EXP5B  both  was  depth.  was  decrease  in  also  TEMP  at  constant  the  the  ppt  during in  with  situ  depth  with  decrease  average  a t 27.2  °C  the same p e r i o d  a significant during  at  statistically  variability  temperature  day-*  four  a  i n Tank B d u r i n g  remained  essentially  was  less  18.1  the  there  with  The  )  16.0  °C  last  time of  0.3  week o f  value.  (SD=0.92)  The during  Regime  Inflow variable  nitrate  with  illustrated inflow  t o both  time,  nitrate to  remaining  period.,  61  periods  N 1-*  (SD=7.70) and  , there  greater The  tide.;  to  c f EXP5A a v e r a g e d 21.9  the f a c t  As d e s c r i b e d  uM  to  14.3  N 1~*  but  the 20  trend  first  uM  1-*  N  N 1~*  (SD=1.24)  during  the t h i r d  sampling this  cn a d a i l y  in  three  concentrations uM  extremely  (av S D = 8 . 8 0 ) .  a decreasing  N03  i n C h a p t e r 5,  partially  uM  for  than  similar  N 1~*  that the  the i n f l o w c o n c e n t r a t i o n  designed  was  inflow  inflow concentrations  attributed  16.5  concentration  levels  three  s y s t e m s was  averaging  i n Figure  increasing  in  °C )  °C ,  there  and  19.8  experiments.  Nutrient  low  (0.6  FB  between  (r=.994),  15,0  (FR=0.50 day-*  salinity  day-*  averaged  temperature averaged  temperature  although  i n TEMP w i t h  day-* Ffl and  the c o r r e l a t i o n  small  (t=1,41) a v e r a g e d (CVf6.1%)*  The  high  but  0.10  °C a t t h e 0.25  FB.  EXP5A, and  the  As the  weeks,  during  the  during  (SD=2.91), 14.1 respectively.  week c o u l d be  time c o i n c i d e d  ;  the uM The  partly  with  low  c a u s e d an  underestimation  basis.  algorithm  compensate f o r the  An  discrepancy.  A  was tidal  ratio  was c a l c u l a t e d  that  the  the  based  on t h e amount  t i d a l h e i g h t was  amount  o f time  above t h e h e i g h t a t s a m p l i n g  p e r day t h a t  at  nitrate  c o n c e n t r a t i o n a t sampling inflow  indicated averaged  time;  the t i d a l  height  daily  sampling  nitrate  t h a t the  when  the  time  nitrate  f o r t=1,41 i n c r e a s e d  period  temporal  value  t o 19.0 uH  effect  the  increase  in  nitrate  varied  e s t i m a t e s of average  day-  recovery  1  the depletion  flushing  rate,  of s i g n i f i c a n t  net  ; N  uptake  of  nitrate  l -  1  and  21.9  uH  N  was  12.0  Nitrate  d e p l e t i o n d i d not occur  based uH N l -  ) and s i g n i f i c a n t apparent  N  l  upon s a m p l i n g to  -  was  account  with  tidal  the  actual  on  o c c u r r e d by Day  there  *as  Days  on 1  by  no  for  the primary  the  until  the  tidal  three  in  spite  of  community  the i n s i t u  1  Tank  The t a n k  -  ,  14. 1  periods; the  N 1-* and 24.1  c o n c e n t r a t i o n s (>2 uK N l 13.  in situ  correction  Day 7 i n  5 at  experiment.  a v e r a g i n g S.3 uH N l "  , 16.3 uM  12 and  in  o f the  1  factor,  uH N l B  ) of  1  .  (Ffi=1.0 nitrate  was v e r y c l e a r a t  t i m e and much o f t h e p h y t o p l a n k t o n s t o c k had  benthos,  utl  included  of n i t r a t e  1~*  were  this  results  underestimated  {N03N)  TNQ3N,  were  The  N 1-* f o r t h e same t i m e  N03 f o r the r e m a i n d e r  values  - 1  revised  o f c h a n g e i n N03  and  corresponding  day  a  16.6  of the t i d e  still  i n c r e a s e d during the experiment, uM  t o the  the e x p e r i m e n t a l p e r i o d . , So t h e r e v i s e d  probably  Tank . A,  0.25  The  TN03  the  inflow n i t r a t e concentration.  In the  during  of  time t o  c o n c e n t r a t i o n with t i d a l h e i g h t ,  s i n c e t h e t i d a l a m p l i t u d e and r a t e height  estimate  TN03(IN).,  day  below  was t h e n a p p l i e d to  t a k e n i n t o a c c o u n t . , However, no f a c t o r for  h e i g h t was  this factor  concentration,  inflow  o f time d u r i n g t h e  circulating  sunk  to  the  pumps. , However  1  70  Skeletonema costatum in  Experiment  (<1  8  uM  1~*  ) during the  Standing  dominant urea  ph y t o p l a n k t e r .  concentrations  phytoplankton  blooms  T a n k s A and  were  E  (35-40 ug C h i a 1-*  magnitude  i n Tank A and  stabilized  by  Day  coincident  ( F i g u r e 65) ).-,  0.25  day-*  doubled  t o 0.5  day-*  ug C h i a 1~  21  Days  19  value  t o 22.  phytoplankton period large  (FR=0.5G  t o Day  21  to  23.  between t h e  45 28,  the  The  the  low  after  the f l u s h i n g  last  phytoplankton  approximately  17.1  double  during  the remaining  6 days  i n the the  inflow  high i n c i d e n t s o l a r  results  was  increased  to  i n Tank  A f t e r t h e FR  phytoplankton  stock  A, was  stock  lowered  decreased  from  increase i n of  the  the  second the from  radiation  Days  was  conditions for  the  1-*  steady-state  levels  t o 0.10  from  a 2 day  lag  productivity  i n c r e a s e i n phytoplanktcit stock,  period  days  concentration  i n d i c a t e d that there forcing  level  c o r r e l a t e d with  nitrate  Day  rate  the  large  significantly  by  three  ug C h i a  The  ) was  apparent  the  at  were o f  characteristic  during  restabilized  more f a v o u r a b l e  .  the  and  with  A, t h e CHLA  period.  28 and  a 1-*  14,  I n Tank  first  resultant  Chi  Day  was  -  The  p e r i o d . ,.  and  1  day *  o f - t h e second ug  on  p o s i t i v e trend  21  end  the stock  Day  and  FR  This  during  12 i n Tank B. ug C h i a 1~*  a t c a . . 8.9  of • t h e  ca.,  were  Stock  minimum i n CHLA f o l l o w i n g n u t r i e n t d e p l e t i o n was 9  As  experiments.  d e p l e t i o n i n both  a similar  the  Dynamics  initial  nutrient  still  4, i n f l o w ammonia and  Phytoplankton  The  was  and had  by  reached  day-* on  to concentrations  the  Day  between  71  15-20 end  ug  C h i a 1-*  of the experiment  tank  by  the  phytoplankton  stock reached  similar  as  A.  Tank  The  t o Tank A,  the  maximum station  levels which  phytoplankton  ug C h i a h »  standing during  cf  the  the  stock  the  primary  significant and T a b l e  day-*  (t>6),  was  stock  doubled  by  ug C h i a 1-* lower  displayed  a  19-22  as  high  N03  Day  some  series  at FB  day-*  a similar  There  depth  and  at  the  of  damped of ca.  period.,  The  c o n c e n t r a t i o n i n Tank B  (t=42,49).  i n c r e a s e i n CHLA w i t h  Ffi  SB  reason.  bloom, d e c r e a s i n g t o a l e v e l 1.0  and 23  , except  for  the  Days  twice  i n c r e a s e i n incoming  of the  remained  0.5  t h e FB  significantly  a t t h e end  stock  the  of c a . , 60  was  p e r i o d i r Tank B  although  phytoplankton  oscillations following this 30  contamination  a s t e a d y - s t a t e v a l u e from  However, a f t e r  time,  reached outflow  t o the  nutrient-depleted  which was  this  due  near-zero values at  protozoans.  During  at  , before approaching  i n both  was  a  small  tanks  (Table  20  21).  Pigment B a t i o s The pigments  concentration  of  CHLB,  CHLC,  are  in Tables  18 and  p i g m e n t s and variable  the this  time A,  0.25  day  i n t h e BA  second higher  pigment  although  I n Tank  the  increase  the c a l c u l a t e d  with  stations., during  summarized  - 1  ratio  FB  19.  ,ratios  generally  after  and  carotenoid A l i the  were not  period,  there  significantly  was  ( F i g u r e 66).., However, by  had a  CHLB  again  increased  during  between occurred  significant the  p e r i o d , CHLB c o n c e n t r a t i o n s were r e d u c e d FB.,  accessory  different  nutrient-depletion  (CT)  end  of  to zero at Period  3,  7 2  contributing  to  CTA  illustrated  ratio i s  between ' the  the  first  CTA  and  decrease  0.1 .from  inflow  exponential  Day  nitrate  Oxygen  16  in  EXP5B i n the  to 22,  oxygen  in situ  oxygen  illustrated  i n Figure  68.,  coincident  with  in  mg  l  -  the  particularly compensate  EA ratio,  except  f o r an  during  for  each i n s j t u  There  In  the  net  was  was  an  increase  with  represented  an  saturation  OXY  was  only  32  was  not  1.00  the  during  six the  tanks,  to low  OXY  maximum  also  a significant Tank  day-*  in  FH  period.  To  inflow  between OXYN and  bottom  an a v e r a g e  a one  station  maximum c f 6.99  of  1-*  160%. higher  much l a r g e r t h a n  for  of  oxygen  calculated for CHLA  mg  significantly l  13.21  -  mg  * on l ~  Day l  In Tank E, t h e a v e r a g e f o r t=1,41 and Tank  A.  was  time l a g f o r CHLA  w h i c h was  oxygen c o n c e n t r a t i o n level  day  during  was  The  with  A,  this  (OXYN) was  correlation  were  difference in A,  the  are  i n Tank  value  1  The  maxima  the  0.10  1~  experiments  bloom, and  In  nag  weeks.  5 was  tanks.  7.26  oxygen i n c r e a s e  the  mg  averaged  first  both  the  particularly  l o w e r , OXYN r e a c h e d  the  variability  station.  (r=.830),  (OXY)  phytoplankton  during  the  (r=.888)• , E x c l u d i n g  Day  During  there  coincident  levels  initial  evident  concentration,  of  tanks.  ),  oxygen between s t a t i o n s i n both  oxygen  f o r both  The  correlation  day-*  a v e r a g e on Day  1  experiment. ,  which  the  (FE=1.00  concentrations  changes  high  and  ratio.,  concentrations.  tanks  • the  high  which was  (av SD=.835) f o r b o t h  13,63  was  67  BA  levels  Inflow  the  i n c r e a s e i n the  Figure  BA r a t i o  s i x weeks o f  exponential ca.  an  the  The  29,  and  an  value  maximum  on  correlation  73  between  OXIN  and CHLA  f o r t h e same  time  Tank B ( r = . 8 1 2 ) , b u t w i t h no t i m e l a g .  swas a l s o  period  high i n  Oxygen s a t u r a t i o n  levels  were s i m i l a r t o Tank A. ,  Primary  Productivity  Results  from  the  primary  productivity  measurements a r e  s u m m a r i z e d i n Taisles 20 and 21., The number c f TIMES in  the analysis  Day  6 a s i n EXP4A a n d EXP4B The  was e l e v e n , r e p r e s e n t e d  gross  rates  illustrated  i n Figure  variability  i n gross productivity  on  4, one day b e f o r e  .initial  A large  in  Tank  1  dy  _ 1  stations  1  PGO  .  r (B)=„98) , decreased respiration  except during rate  The d a t a f o r Day i n E x p e r i m e n t 5. 1  t h e CHLA  and  was a since  were  with  t h e CHLA  l  hr  similar the the (RES)  - 1  PGO  no  daily  was  the  bloom  relatively  15 i s m i s s i n g  a  occurred  differences  variables  experiments on  a  Day  (PGQDY) o f  and t h e  160 ug C 1  The p a t t e r n  productivity  secondary  value  significant  respectively.  gross  i s  occurred  also  EXP5A and EXP5B was  during  are  significant  maximum on 1  f o r any o f t h e p r o d u c t i v i t y during  B  maximum, w h i l e i n Tank B t h e  1  There  h r - * a n d 304 ng C l ~ and PGODY was  A  maximum o f 460 ug C ..Ir' •hr-'  average gross p r o d u c t i v i t y -  Tacks  time  on Day 33, w i t h an e s t i m a t e d  3*83 mg C 1 ~ between  with  was c o i n c i d e n t  secondary  B  day from  However, i n Tank ft, t h e uaximum r a t e  PGO maximum  7.  in  69. . I n b o t h c a s e s t h e r e  o f CHLA.  Day  by e v e r y t h i r d  used  » .  productivity  function  (days)  between  (r(A)=.96; daily  basis  (Figure  70).  The  Constant  during  f o r a l l productivity  both  variables  74  experiments, was  there a marginal  (Figure (PNO) the  a v e r a g i n g c a . , 42  71),  was  The  based  significantly correlation (av  .  ug  61.  which  represented  C  0.84  illustrated The  Maximum ug C  daily  mg C  1~*  of  was  the  standardized ASS  curves  standardized  during  Tank  21  3  The  experiments tanks,  mg  o f CHLA. , The  illustrated  ^he  ug  period  C  During  in  and  C  1-.*  time  are  74  PGOST, to  the  A subsequent  PGOST first  *as r e l a t i v e l y  20.7  to  the  maxima were  also  in  on  Tank  two  time  Day stock  B  was  weeks,  the  constant,  ranging  during the  second  h i g h between t h e  with  76,,  r a t e s of a t l e a s t  day-* ) when t h e s t a n d i n g  variability  the t h i r d  primary  resulting  (FR=0.50 day-* ) and  the c o r r e l a t i o n  the  on  productivity  Figures  secondary  pattern for  During  the  (ug C h l a ) - * h r ~ * . . O n l y  was  rates  unit  gross p r o d u c t i v i t y  10-15  flushing  comparing  (FR=0. 10  different..  PGOST, a l t h o u g h B.  also  f o r Tank B  c h a n g e s i n PCDY w i t h  during Period 2  Period  standardized  for  than  i n t h e two  - 1  and  72).  the  by  bloom,  approached z e r o ,  two-week  The  method  was  r a t e s o f 0.51  were a t t a i n e d i n Tank  -  on  ca.,  (Figure  hr  1  gross p r o d u c t i v i t y  apparent  Day  per  are  phytoplankton  from  -  time  (r>. 97)  (PROD)  during  ug C l  variable  analyzed  -  somewhat  .  tanks  (av £ ( & ) = . 6 5 )  productivity  dy~*  (ug C h l a ) * h r *  level  net  101  with  cn t h e o x y g e n  PHO  PROD r a t e s  and  -  EES  method  and A  in  i n both  radiocarbon  average  initial  39  PGO  based  o n l y i n Tank B  i n F i g u r e 73. ,  productivity  RESST and  with  w i t h PGO  l""* hr *  effect  variables  the  .hr-*, and  difference  g r e a t e r i n Tank The  1~*  productivity  correlated  were  and  net  on  was  r (B) =.47)  dy-*  significant  highly correlated  rate  ;  ug C  was  again  two  tanks  less  in  two-week p e r i o d , EGOST s i g n i f i c a n t l y  75  i n c r e a s e d t o a maximum of c a . B, hr~  while  the a v e r a g e v a l u e s  i n Tank  l  values  in  a until  when  respiration  with  time  hr  ca.  13% f o r Tank  B  76),  variability  rates  in  averaging  at  2 i n both  response  to  c o n c e n t r a t i o n . „• 3 i n spite  from c a .  (ug C h l a ) - * maximum  (see P a r s o n s  ug C  lost  22).  ug  large  during  C  (ug  3 uM N l ~  l  on Day  the  t h e second  ug c  (Ug  corresponding  respiration  rate  Period Chla)  -  .  1  decrease  in  1  in  far  - 1  the  the  linear  -1  was EXpSA  .  magnitude  )  inflow  A close  ( F i g u r e 76)  during  B on Day  Period  examination suggested  increase i n inflow 1~*  27  nitrate  i n Tank a d u r i n g  20 u M N  The  The a s s i m i l a t i o n  i n Tank  i n FJB.  i n Tank B  20 t o c a .  (ASS)  (ug C h l a ) - * h r ~ *  increase  The  C  day-* i n Tank a ,  though  The r a t e a l s o i n c r e a s e d  features.  to r e s p i r a t i o n  rate  t a n k s . , The r a t e i n c r e a s e d the  period  3 i n Tank E.  (ug C h l a ) — * h r  ( 2.5  bloom  During  r e d u c e d t o 0.1  B was l e s s e v e n even  6.5  more v a r i a b l e  the  productivity  6.0  o f the f i v e - f o l d  interesting  during  (Table  period  o f a s s a s a f u n c t i o n of, STATION two  Tank  Pmax's f o u n d i n  (RESST) was  f o r EXP5B d u r i n g  net  were t h e t h e l o w e s t  Period  rate  by damped o s c i l l a t i o n s  i n Tank  greater  in  1  These  the high  the s t a n d a r d i z e d  20% d u r i n g  standardized  (Figure  was  by 7 5 % w h i l e  by o n l y  characterized  -  t o a low r a t e o f 1.5 ug  a f t e r t h e PR was  RESST i n c r e a s e d  The  39.  p r o d u c t i v i t y was  , t h e same v a l u e  - 1  period.,  increased  tban  ( F i g u r e 75) , e s p e c i a l l y  p e r i o d o f EXP5A, RES ST d e c r e a s e d Chla)-*  Day  8 ug C  a S k e l e t o ne ma c p s t a t a m bloom  c a . „. 45% o f t h e g r o s s  compared  at ca.  hr  - 1  197 33).=  The s t a n d a r d i z e d EXP5A  (ug C h l a )  on  both t a n k s a r e g r e a t e r  Takahashi,  during  remained  the increase  Tokyo Bay, a l s o d u r i n g and  20 ug C  by . Day  N03 24  76  failed  .to  phytoplankton limited the  by  alleviate  the  by Day 24.  However, t h e s y s t e m had become  Day  nutrient-limited  27, i n d i c a t e d  t y the high  s u r f a c e compared w i t h t h e b o t t o m  phytoplankton  stock  appeared  outlined gross  i n Chapter  component 5 was used  productivity  analysis to  represented  and  productivity  16$ l o w e r  the r a t i o o f e x u d a t i o n  evaluate  the  The  e s t i m a t e s o f BPGO  (assimilation:gross respectively,  i n T a n k B.  productivity  or  only  was  probably  data  the times  t o 0,50,  then  ESTPGO c l o s e l y a p p r o x i m a t e d  The  of  estimated  1.00 i n Tank A  (EPGO)  of t h e gross in was  experimental  of  EXP5B,  was s u s p e c t . and APGO  and  period  0.36  the  EXP5B,  the  channeled  proportion  exudation  slightly  was 15%, a b o u t  i n EXP5A,  value each  values  of  1.00.  represented  with c a . , 40% a t t r i b u t e d t o  largest  through  component d e c r e a s e d fraction  and e x u d a t i o n  productivity  T h e r e f o r e EPGO  experiment  expected  on  (t=11)  hand, i f t h e  on the t o t a l  Summarizing,, i n EXP5ft r e s p i r a t i o n  respiration  proportion  C.15  On t h e o t h e r  and APGO were based  assimilation  t h e one  productivity)  were  for the total  BPGO  productivity  Model  to  case  when EXC was measured ( t = 5 ) .  closer  assimilation;  i n the  ( r e s p i r a t i o n : gross  for  30%  the  r e g a r d l e s s o f whether t h e e s t i m a t e s were b a s e d  the  ca.  23. ,  productivity  productivity)  33,  r e s p i r a t i o n and  was 6% h i g h e r t h a n  t o gross  Day  similar  ty a s s i n i l a t i o n ,  (ESTPGO)  than e x p e c t e d  By  rate at  Productivity  e x u d a t i o n , , T h e r e s u l t s a r e shown i n T a b l e gross  light-  n i t r a t e - l i m i t e d again..  E s t i m a t i o n of Parameters f o r j Primary ft p r o d u c t i v i t y  of the  assimilation  station.  t o become  state  half  of  ( ca.  to c a .  gross  50% ) , the  35%  and t h e  t h e v a l u e i n EXP5A.,  77  Values initial  f o r ALPHAC  slope  relationship, in  Chapter  C  (ug  (ly/min)—  5.,, I n  **C  hr~*  - 1  during  1  14.0  15.7  C  inflowing  value  very  low  size  1  and  e v o l u t i o n of the  Figures  85  representative approximately EXP5A.  The  are  of one  90  week a f t e r e a c h  corresponding  80),  the  micron  results  12.  initial  maximum v o l u m e - o f size  decreased Day  the  on Day  6;  with  with bloom  the  Chla)-  the  ug and hr-  1  1  value f o r  compared  f o r EXP5B.  high.  on  The  tanks  with lowest  when  the  The  corresponding  gross  productivity  Community  communities i n terms of  i n F i g u r e s 79  of  change from  t o 84  s i x times  the  a high  i n the  to  maximum  EXP5A  o r days  primary  were  community  flushing  rate i n  constant f l u s h i n g  p e r i o d i n EXP5A  the o n s e t  for  Tank E were a l s o i n c l u d e d  •particles'  maximum volume s h i f t e d By c o m p a r i s o n ,  was  station  average  2 1 i n both  60  as  78.  structure  a s a c o m p a r i s o n of a s y s t e m During  The  f o r E X P 5 E . „ The  the  (PAH)  PA8ZC d a t a  (ug  (ly/min)-*  phytoplankton  illustrated  to  SB  i n Figure  o f the P h y t o p l a n k t o n  composition  77).  s l o p e based  (ALPHAG) a r e i l l u s t r a t e d  The  and  and  bottom C  ( l y / m i n ) -* Cay  light  a maximum of c a .  ug  -  Chla)-* hr-*  was  11.7  (figure  ASS  the  (ug C h l a ) - * h r *  e s t i m a t e s of. t h e i n i t i a l  Composition  of  ( l y / m i n ) - * ) , the  versus  the  at  -  ALPHAC o c c u r r e d on  N03  from  (ly/min) *  Period 3  ug C (ug  e s t i m a t e s of  productivity  EXP5A, ALPHAC r e a c h e d  t o an a v e r a g e  EXP5A was ug  the  (ug C h l a ) - * h r - *  were a l s o e s t i m a t e d  Chla)  decreased  in  ( ug C  was  found  rate.  ( F i g u r e s 79 at  the  of n u t r i e n t - d e p l e t i o n ,  the  14.3  u diameter  volume i n Tank B  was  size  and 22.6 the by  evident  78  at  the  same d i a m e t e r  and  86).  The  later  o f 22.6  u on  timing of the  9 f o l l o w e d t h e same t r e n d a s  Days 6,  maximum  the  9 and  12  particle  phytoplankton  ( F i g u r e s 85  volume on  stock  Day  measured  as  CHLA. .... During A  shifted  9.0  u and  Period 2  to a bimodal 18.5  u  phytoplankton although u  by  (Days 15-28), t h e  sizes  in  81  retained  of the  a  and  82).  unimodal  maximum volume  maximum volume i n c r e a s e d a g a i n t o 14.3  u by  During  concentration maximum  87  Period  significantly  at  by Day  39  ( F i g u r e s 46  Skeletonema costatum o f FXP5B e v e n on  d e c i p e n s J7J. and  diatom  species  Nitzschia  composition  Chaetoceros after also  species  was  (compressum?)  t h e i n i t i a l bloom present  Tank ft had  a  and  in significant  at the  present  not  the  decreased  the  end  during  of the  at a  B  were  although 1.0  day-*  EXP5A was  quite  in significant  the  numbers,  and  Nitzschia  Throughout the with  A  dominant p h y t o p l a n k t e r  Thalassiothrix  much h i g h e r d i v e r s i t y ,  Tank  was  dominant p h y t o p l a n k t e r .  was  numbers.  in  initially  c l o s t e r i u m (?) ,  FR  Although  with  t  phytoplankton  apparent  phytoplankton  volume  dominant p h y t o p l a n k t e r f o r the  some B h i z o s o l e n i a were a l s o  costatum  phytoplankton  stock  main  Skeletonema  89  the  irhe o t h e r two  EXP5B.  (Figures  was  minimum,  from  39  11.3 the  distribution  47).  the  of  12 when t h e s y s t e m  and  different  Day  the  to  diameter  Day  nutrients  The  cell  the  of  .,  shifted  t h e m a g n i t u d e had and  Tank  distribution,  was  depleted  Chaetoceros  size  11.3u, a l t h o u g h  in  However,  size  had  3 i n EXP5A, t h e  r e t u r n e d t o a unimodal  volume  duration  and  composition  maximum v o l u m e s a t  The  90).  (Figures  with  88).  and  24  (Figures  EXP5B  the diameter  Day  distribution  size  were  experiment.  more s p e c i e s o f  diatoms  79  and  a  species  variety list,  unicellular and  of  nano-flagellates.,  also  included  Chaetoceros  { Oxhyrris  sp.. ).  composition,  as well  affected  the  by  •contaminated* experiment. ,  the  ,  cryptomomads be  of  the  system  delicatula  ,  ( Cryptomonas s p .  noted  a s c e l l numbers,  primary  t h e end o f EXP5A, t h e  fihizpsclenia  I t should  presence  By  that  the  A would have  predatory  ciliates  the l a s t  , )  species  i n Tank  during  a  been which  weejk o f t h e  80  CHAPTEB 7 .  TWO-STAG! CO H U M OOPS COLT OSES OF PLANKTONIC  HEfiBIVOBQOS FOOD CHAINS: G.BQ1TH OF THE HERBIVOEES  Experimental  Design  Growth o f t h e s e s s i l e gjgas 5)  •,  was i n v e s t i g a t e d  using the outflow  continuous  source  herbivore tanks.  *planktonic*  i n a two-stage c u l t u r e  from e i t h e r of  Four  period  differed  in  flushing  rate  )  to  of the  primary  phytoplankton  the  fron  primary  Days  source o f the inflow  compare  various  (flagellate,  diatom,  phytoplankton  and  types  or  other  of  mixed) growth  Crassostrea  system  (Figure  A  or  B  as  a  oxygen  to  the four  (EXFI t o EXPIV) o f one  18 t o 50 d u r i n g t h e p o s t -  tanks..  o f the herbivore tanks  tank  and  oyster experiments  week d u r a t i o n were c o n d u c t e d bloom  herbivore,  The (Tack  four  experiments  A o r B) and i n t h e  ( F i = 1.0 d a y *  o r 2.0 d a y  -  phytoplankton  - 1  communities  and v a r i o u s c o n c e n t r a t i o n s o f variables.  The  experimental  d e s i g n i s shown i n T a b l e 16. As  described  oysters  juvenile groups:  with  oysters very s m a l l  and v e r y l a r g e  attached range •small*  2, Tank 4 was t h e c o n t r o l  and Tanks 1, 2 and 3 were s t o c k e d  respectively,  mm)  i n Chapter  8 o y s t e r s per a r t i f i c i a l were s e l e c t e d  and then  (35-45 mm), s m a l l  2, 4 and 8 c u l t c h  cultch.,  divided  (45-55  nm),  96  into  healthy  four  large  size  (55-65  (65-75 mm). , The 8 * v e r y s m a l l * o y s t e r s were  t o C u l t c h #1, w h i l e t h e 32  were  with  w i t h no  randomly  chosen  oysters  f o r placement  c u l t c h .( #2 t o #5).. A s i m i l a r  in  the  45-55  mm  on one o f t h e f o u r  procedure  was  used  to  81  produce large*  the  four  cultch  (#10  Therefore some the  extent  "large  1  cultch  t o #12),  giving  ca..  held 24  growth  the r e g u i r e d  as a f u n c t i o n  of t h e i r  density  the t h r e e * very twelve  the p r o d u c t i o n o f o y s t e r s c o u l d  i n the o u t f l o w from  hours.  Initial  variables  be  and  for  one  weight)  week  size  arid t h e  chlorophyll a concentration, once  a  day  experiment,  at  ca.  Results F i g u r e s 91  and  hours  were a g a i n weighed from  for  and  p r o d u c t i o n of  the  oyster  in situ  T a b l e s 24 t o 26.,  derivation  four  (P=.001)  conducted level,  monitored end  of  each  measured and  the  are  Appendix  of a d d i t i o n a l  summarized  in  contains  the  2  variables pertinent  to  herbivores.,  temperatures  experiments are i l l u s t r a t e d significant  the  meat  the herbivore tanks.,  experiments  Environmental Conditions during The  and  oyster  weight,  were  fit  tank f o r  oxygen  ammonia  PST. ,  cultch  the  experiments  of  Results  t o 95 and  description  the  total  temperature,  urea  f e c e s and p s e u d o f e c e s c o l l e c t e d  Experimental  oyster  The  1200  the oysters  depth,  to  i n each  the a p p r o p r i a t e p r i m a r y  width,  periods.,  cultch.  examined  measurements were t a k e n f o r  (length,  weight and s h e l l  the  and  e x p e r i m e n t s . .., B e f o r e e a c h h e r b i v o r e e x p e r i m e n t ,  were  the  (#6 t o #9)  difference experiments,  in  t h e O y s t e r Growth during  the  i n F i g u r e 91. temperature  there  was  i n TEMP between e x p e r i m e n t s  a  Experiments  oyster  growth  A l t h o u g h t h e r e was  no  between t a n k s i n any  of  significant (Table  24) .  difference In  Experiment  82  3,  the temperature °C  (23.3  ), r e s u l t i n g  source inflow Tank  B  d u r i n g t h e week a v e r a g e d  1.0  1 was  t h e l o w e s t and  d a y - * ., The  The  temperatures  19.5  °C  in  experiments. Experiment  The 3  indicating  in  low  was  Figure  in  Experiment  temperature  during  2 and 4  the from  3  and was  i n Experiment  t h e one  92,  week  averaged  use  averaged  30$  in  coincident  increased ,1  tank,  and  with  the  the  mentioned  of  lower  period.  20.2  °G  and  phytoplankton and  latter  between  part  o f t h e s t o c k was in situ  Experiments  t o 58% a t the  i n Chapter  In a l l  3.  Since  be  6.,  day-*  but  the  control  concentration,  t o the benthos i n pumps.,  The  day * -  flushing  loss  flushing rate  in  t h e r e were nc h e r b i v o r e s i n the  h i g h e r compared  which had a s i g n i f i c a n t  lest  4 a t t h e 2.0  1.0  the  of  circulating 2 and  of  contamination  than the i n f l o w  these percentage l o s s e s o f  would  inflow within  i n f l o w CHLA d u r i n g  a proportion  Experiments  the  considerably  significantly  the  the  than  the phytoplankton c o n c e n t r a t i o n i n the  of  sinking  °C a v e r a g e  Tank .A by p r o t o z o a n s , a s  (4) was  control  higher  of  f o r the e t h e r t h r e e experiments)  Experiments  varied  f o u r experiments*  and  value  respectively.  concentrations  rate  18.5  highest  temperature  significantly  most v a r i a b l e  As i l l u s t r a t e d  spite  f a c t o r s . , The  rate of the herbivore tanks  only  tank  two  from Tank A was  (the s o u r c e i n f l o w  the f l u s h i n g  primary  from  the  phytoplankton  due  to  to the other h e r b i v o r e tanks  variable  grazing  rate  depending  on  number o f o y s t e r s p e r t a n k . In  sinking  view  of the  rates  phytoplankton  lack of d i r e c t  within  (STKUP) was  each  measurements o f p h y t o p l a n k t o n  herbivore  e s t i m a t e d as  tank,  the  the  uptake  difference  of  between  83  the  inflow  and  in situ  maximum e s t i m a t e . decreased period  fairly  during  constant  phytoplankton  herbivore  tanks  the l a s t  difference  I-*  and  uptake and  there  (?=.738).  were no  as  phytoplankton Experiments  ug C h i a is  with  stock  one  week  phytoplankton  low  In EXPI, in  all.  there  the as  source  was  4  , more t h a n  , 24.5  l  1~*  was  a  three  significant  had  since  flagellate  flushing in  expressed  as an  average  the uptake  Experiments  2 and  tanks.  4 was The  in  the  ratio  between  1,  the  rate  three  phytoplankton  2 and  rate  (STKUPE)  ,  mg  a  Chi  1.,  Since  tanks,  to  a  only i n  difference i n  STKUP  s t o c k l e v e l s i n Tank 3 than  4  the  3 were g r a z e d  oyster  there a s i g n i f i c a n t  a food  when  i n Experiment  II and  only  experiments  and  uptake  ug  (Table  as  v a l u e s o f 8.4  t h e h i g h e s t d e n s i t y o f o y s t e r s were l e s s E X P I I and  was  r a t e s i n Experiments  t h e stoc-k c o n c e n t r a t i o n i n E x p e r i m e n t s zero  it  sith  , 7.8  1 to 4  community  Experiment  similar  double  approaching  ug C h i a 1~»  i n Experiments  excluded*  the  high  2 and  the  increased inflow concentration  d i f f e r e n c e s i n STKUP  However,  twice  The  of  EXPIV.  remained  tank,  ug C h i a l ~  i f EXPIH  source,  both  standing  of the experiment. .  24.8  experiment  between  provides a  to 3 during the  the  EXPII  of the  the. c o n t r o l  growth  level  and  (P=.QGO) i n STKUP between t h e f o u r e x p e r i m e n t s ,  24) ; a l t h o u g h  dy-»  1  and  stock l e v e l s  v a l u e s a v e r a g i n g 22.3 a  stock  phytoplankton  Tanks  in  i n spite half  Excluding  were  in  i n a l l four experiments,  situ  Chi  in situ  exponentially  remained in  The  phytoplankton  2 ug C h i a 1  _ 1  with in  EXPIV. , of  summarized i n T a b l e  chlorophyll 24,  b to chlorophyll a  s i n c e i t g i v e s some  (B&) i s a l s o  indication  of  the  84  species  composition  experiments, were  of  phytoplankton  t h e BA o f t h e p h y t o p l a n k t o n  significantly  similar  the  value  higher  to  preferentially  the  graze  than  the  inflow.„,  apparent  between  oysters  the l a r g e diatoms,  densities,  the  herbivore  particularly  inflow  oxygen l e v e l s  oxygen inflow  i n Figure  uptake  and i n s i t u  93.  3.77  mg  1-*  ,  respectively. the:!  three  experiment  to  Furthermore,  the  tank c o n t a i n e d contrast  to  significant the  least  i n EXPI  -*  )  in  EXPI  1—  uptake  the  .,  1  was a l s o  low t o h i g h  other  experiments  are  s t o c k , the  and  the net  uptake o f oxygen  the c o n t r o l  3.85  tank,  »as  mg 1~» and 5.23 mg 1-*  3, the oxygen c o n c e n t r a t i o n  above  uptake  during  the  the l a s t  inflow  density  three  cf  concentration.  oysters.  experiments,  o f oxygen  was e x p r e s s e d  contrast t o the r e s u l t s i t i s apparent values  This  i n which  there  Tanks  1  this  is in was a to  3.  a s a r a t e , OXYDPfi was i n EXPII  f o r STKUP8. that  in,  h a l f o f the  i n Tank 3 was t h e l e a s t a l t h o u g h  (p=.006) i n 0XYUP between  with  the  light  d i f f e r e n c e between t h e  ( 640 mg d y - i ) a n d g r e a t e s t  (0.17),  trend  experiments  The a v e r a g e  increased  the highest the  as  Experiment  levels  OXYUPR:STKUPR r a t i o , for,  mg  tanks  increase  When  between  1 t o 4, e x c l u d i n g  5.45  oyster  from  high  the phytoplankton  concentrations.  During  (i.e. a  A similar  tanks  As w i t h  calculated  (OxYUP) i n E x p e r i m e n t s  to  l e a v i n g an e n v i r o n m e n t  f o r the four oyster  varied  was  appear  i n EXPI and EXPIV. ,  Oxygen c o n c e n t r a t i o n s illustrated  tanks  c o n t r o l which r e t a i n e d a  The  temperature, low n u t r i e n t s y s t e m ) .  In a l l  i n the h e r b i v o r e  c o n d u c i v e t o the growth o f n a n o - f l a g e l l a t e s and  stock.  this  (1852 mg  dy  By c o m p a r i n g the ratio  was  least  o f 0.22 f o r E X P I I and EXPIV and a  85  large  maximum r a t i o  results  increase  and  between  in  existed  USEA c o n c e n t r a t i o n s  of  the  The  means  variables  cultch  weight) (n=8)  the t o t a l  and  oyster  at  weight  cm;  by was  cm  (12%).  experiment  representing  NH3  steady and  no s i g n i f i c a n t  in  terms  (EXP), d e n s i t y  deviations  width, depth, t o t a l  the  for  NH3  increase  Factors of  the  three  (DENS) and and  of  size  illustrated  d e p t h (D)=  1.7  the  the  cm;  twelve  weights,  total weight  weight  total of  (181)  weight i n c r e a s e d the  total  by  50%  weight,  the  oyster  experiment  (SGTT) =  for  5.7  cm;  12.8  increased  g,  was  g;  g. . at t h e  depth  t c 19,2 which  and  averaged  (BGTS) - 8.8  and  growth  meat w e i g h t  (N=96), w e r e : l e n g t h (L)=  g; s h e l l  cm  six  each .herbivore  d i m e n s i o n s and  of o y s t e r s  4.0  of  for weight,  each  beginning  initial  (flGT«)=  32%  a  phytcplankton  i n T a b l e 25 and T a b l e 26  (7%) , t h e w i d t h by 0.6 The  uM  difference  that  e n d o f a l l f o u r e x p e r i m e n t s , t h e a v e r a g e l e n g t h had 0.4  with  , 1.4  u£i Nii3  no ' s i g n i f i c a n t  growth  standard  population  width(W)= 3.2  There  experiments,  indicate  uptake  were c a l c u l a t e d  The  1--* , 0 . 8  d i d not  95.  (length,  (Table 25).  meat  and  these  either.,  a r e summarized  i n F i g u r e s 94  the  a F u n c t i o n o f t h e E x p e r i mental  experimental factors,  shell  NH3  the h e r b i v o r e s .  Eesults  of  and t h e r e s u l t s  between  O y s t e r Growth as  product cf oysters,  , , T h e r e was  1  Furthermore,  with temperature.  any  ufl NH3  uu N H 3 1~  tanks e i t h e r ,  (SIZE))  in  r a t e s o f 1.9  by  EXPIII. ,  the primary e x c r e t o r y  0.6  excretion  for  correlated  significantly  production  state  0.49  are p o s i t i v e l y  ammonia,  l"-»  of  by  by 0.2  w i t h HGTM the  same  86  proportion The  as the i n i t i a l n e t and p e r c e n t  calculated averages three  summarized  i n Table  during  EXPII  each experiment  represent  the  experimental  (NETL, NETW  (P>.001)  between  NETD)  experiments. cm  were  NETH and NETD r e s p e c t i v e l y ;  4 0%  period  of the t o t a l  (four  weeks)  f o r a l l three  values  3.2% week-* f o r PEEL, 7.5% week-* f o r PEHW a n d 4.7 %  population  growth r a t e s o f l e s s than  1.2% p e r week.  T h e r e were a l s o s i g n i f i c a n t  in  the net increases terms  of  EXPIV, with and  1.4  basis  values  g f o r NETHS.  for total  (PEEBfi-15.4%) of  growth  The  %  largest  weight a l s o  increase weight  in  netincreases  EXPIV,  occurred  measured  occurred i n  i nthe total i n EXPII  with  experiments  f o r NETKT, 0.7 g f o r  Large n e t i n c r e a s e s  and s h e l l  was  occurred  v a r i a b l e s which were  a v e r a g e s o f 2.1 g / o y s t e r  of  represents  d i f f e r e n c e s between  i n t h e growth  weight..  meat w e i g h t a n d s h e l l the  this  The lowest average n e t i n c r e a s e s i n t h e l i n e a r  dimensions o f the oyster  in  *  for  I n terms o f  weekT* f o r PESD.  increases  increase  variables. of  percent  for a l l  and  o f 0.17 cm, 0.24 cm and 0.08  more t h a n  were  and t h e  Average n e t i n c r e a s e s  f o r NETL,  these values entire  26.  different  average i n c r e a s e s  attained  during  measurements o f growth  significantly  Maximum  increases  f o r a l l s i x growth v a r i a b l e s f o r each o y s t e r  dimensional  were  measurements.  ,  NETHM weight,  which  on  i n growth, were t h e l a r g e s t a v e r a g e  (PE£WT=13.51 weight  week * -  ),  meat  weight  (PEBHS= 12. 6%) .'• , A s i m i l a r p a t t e r n  a p p a r e n t i n EXPIV  and EXPI, w i t h  slightly  lower  The % growth r a t e was c a l c u l a t e d f o r e a c h o y s t e r as the net increase during the experimental p e r i o d d i v i d e d by t h e v a l u e a t the b e g i n n i n g o f t h a t , e x p e r i m e n t . 1  87  percentage increases. increase  by  the  total  t o ca.-, 0 . 1 7 f r o m  dropped was  in  During EXPIII, there  low compared  weight  with the o r i g i n a l  The  increase  dependent  each  corresponding medium  i s broken  density  in  Tank 3.  There  linear  dimensions  particularly linear,  was g e n e r a l l y with  a  an  The  results  similar  definite  EXPIII  were found  medium d e n s i t y terms  significant  oysters low  by  oyster  density,  i n Tank  1,  in  increasing  growth with  DENS=2  the  density  pattern.,  tank  in  density  variables, total  w i t h an i n c r e a s i n g d e n s i t y i n t h e medium d e n s i t y  persisted  only  although  i n the  case  of  and  (2) had t h e l o w e s t  decrease  tank,  case  The h i g h e s t  i n the high  fora l l in  DENS=1 to the  density i n  average of  net  oysters,  However t h e d i f f e r e n c e s i n t h e  f o r EXPIII  o f t h e weight  density  growth  the  (SIZE)  d i m e n s i o n s between DEWS ware n o t s i g n i f i c a n t  EXPII.,  weight  94,  decrease  i n EXPI and E X P I I . ,  Finally,  i n EXPIV.,,  Tank 2 and DENS=3 t o t h e h i g h e s t  percentage basis, except i n the  In  of 0 . 3 2 ,hut  i n growth a r e i l l u s t r a t e d a s  down  to t h e lowest d e n s i t y  8.1%  EXPII. , I t  0.34.  (DENS) and s i z e  variables.. In Figure  experiment  and  to  occurred  an  o f NETWM:NETWT  HGTH:WGTT r a t i o  i n s h e l l weight  n e t and p e r c e n t i n c r e a s e s  only  ratio  EXPI  had r e t u r n e d  a f u n c t i o n of the f a c t o r s density six  the  the 0 . 3 7 d u r i n g  t h e e n d o f EXPIV, t h e r a t i o  thesmallest  and  sas  depth  EXPIV  linear  on a n e t o r  f o r EXPI  and  d i d not f o l l o w a growth  rates  in  t a n k , w h i l e i n EXPIV, t h e rates. there  weight,  was  generally  meat w e i g h t  a  and s h e l l  o f o y s t e r s . , However i n E X P I I ,  t a n k grew  more than t h o s e  i n the  on a p e r c e n t a g e b a s i s , t h i s , t r e n d of  NETWM. ,  When  considering  the  i n w e i g h t on a p e r c e n t a g e b a s i s , t h e s i g n i f i c a n c e l e v e l s  88  were l e s s  and  t h e r e was  no s i g n i f i c a n t  DENS i n any  o f the e x p e r i m e n t s .  of  o f the secondary system  a lew  of by  FB  phytoplankton  was  I t appears  limiting  and  i n the  growth..  high  density  total  weight  The  effect  and EXPIV  a significant and  more  SIZE  between  experiments  consideration  (Figure 95).  was  generally  particularly there  effect terms had  i n EXPI and  (Table o f the  26).,  weight  percentage  significant  SIZE  EXPII..,  on  differences  experiments., i n EXPIII.„  was  growth r a t e s . the  effect  covariates  In f a c t  cf oysters  the  weight  differed  variables  under  dimensions,  there  P.EB.L w i t h i n c r e a s i n g  size,  the  other  hand,  although  i n w i d t h between t h e s i z e no  larger  significant  size  3 and  i n a l l experiments, was  weight  was  of  linear  PEBD., I n  H  oysters  although  r e v e r s e d . ,.: T h e r e  upon PEBwT and  Growth i n meat  the o n l y  similar.  a l s o t r u e o f NETD and  trend  of size  cn  was  variables, the  basis,  linear  This  in  difference in  between s h e l l  of l i n e a r  and  i n some e x p e r i m e n t s , t h e r e  the h i g h e s t net  four  depending  a d e c r e a s e i n NET!  were s i g n i f i c a n t  oysters  pattern  terms  the  the type o f p h y t o p l a n k t o n  upon growth r a t e s  In  only  limited  v a l u e s o f the  to EXPII.  was  the  were  as a f u n c t i o n o f DENS, were of  combination  tanks i n EXPII,  (DENS=3)  (STKUP, OXYUP and TEHP) were s i m i l a r  and  the  i n growth i n E X p i i l ;  tank  community., W i t h i n e x p e r i m e n t s ,  with  an i n c r e a s e  g r o w t h between DENS a l t h o u g h t h e FB  between E X P I I  PEBWfl  inflow concentration  However, i n EXPIV t h e r e was  difference  that  in  low  d o u b l i n g the flow r a t e i n the o y s t e r  oysters  a  increase  PEBWS  in  on  was  a  a l l  not a f u n c t i o n  of  89  CHAPTEB 8./  ANALYSIS OF  Estimation of  the  THE  BESULTS USING J  SIMULATION  P h y s i o l o g i c a l Parameters d u r i n g the  MODEL  Continuous  Cultures  In  order  primary  to estimate  p r o d u c t i v i t y model o f t h i s  e x p e r i m e n t s cn light,  photosynthetic  temperature  simultaneously the  surface  days  one C  with  the  or  20  p r i m a r y tank Primary by  the  uptake c f  period each  from  1030  compartment  sclarimeter outlined  and  t o 1430 of  the  i n Chapter  the PAB  2.  v  period  was The  function  from  consecutive  7) , and  was  ( 14 °C  of  conducted  were t a k e n  four  (PAB)  a  s e r i e s of  were  (Days 3 t o  repeated  the  on  each  examined , 16  f o r the  °C  three  at  , 18  °  highest  of n u t r i e n t d e p l e t i o n  phytoplankton  incubator,  d i f f e r e n c e i n SB  a  for  in  12) for  during  as  cn  temperatures  neutral density  out  )  radioactive  water-cooled  carried  day-*  was  a short  samples  intensity  initial  (Days 10 t o  i r r a d i a n c e using  significant  the  productivity  constructed  were  light  values  (nitrate)  blccm p e r i o d  ) . , T h i s process  temperatures during  measured  1*0  four experimental °C  productivity  Phytoplankton  (F.EU =  initial  parametric  ecosystea,  nutrients  EXP5.  p r o d u c t i v i t y versus of the  the  and  o f Tank B  during  day,  appropriate  carbct at  eight  filters.  clear,  in  All  between d a y s d u r i n g  incubator  was  c a l c u l a t e d as techniques  for  The  a  levels the  b r i g h t days and  h o u r s P.S.T.  samples  the  total  measured 0.50 the  was  specially of  natural  experiments there  was  no  incubation radiation in with  of t h i s  value  measurement  the as of  90  primary  productivity  and  standing  stock  are  also described  there. , The in  r e s u l t s of the  Figure  96.  normalized pass  the  approximates  net  Eppley  conditions,  and  the  between P the  and  6  PAR  asymptote  estimation functions The between  of  the  data  curves  parameters  of  s e t s of  with  was  a  nutrient  temperature.  apparent  not  the  method  indicated  f o r a l l four  were  not  (McAllister et a l .  increased  points  t h e c u r v e s do  f o r both  intensity  were  at  the  experimental  included  various  in  the  mathematical  tested. two  common p a r a m e t e r s  primary  ( ug C  the  these  light  illustrated  radiocarbon  productivity The  are  productivity  as e x p e c t e d ,  Sloan,1965).  experimental  temperatures;  of  primary  since  particulate  and  experiments  t h e bloom p e r i o d , p h o t o - i n h i b i t i o n  highest  ALPHA  of  origin  hyperbolic relation  During  values  I*  i n t e r m s o f CHLA, a n d  through  ,1964;  All  *F v e r s u s  light  productivity  (ug C h l a )  - 1  saturation  ) , t h e asymptote  or  were  for  estimated  hr  i n the mathematical  and  - 1  light,  (ly/min)-*  curve,  and  IMAx  photosynthetic rate the  h y p e r b o l i c models were used  seven  'P  namely t h e  relationship initial  ) of the l i n e a r ( ug  C  portion  (ug C h l a ) - *  at optimal  slope  hr  irradiation,  versus I* experiments.  to d e s c r i b e the  - 4  Two  data: the h y p e r b o l i c  tangent  P and P  6  6  (I) =  PMAX * T ANB( ALPHA*X (I) /PM AX)  Smith's function  -  R  6  ( S m i t h , 1936)  (I) = PMAX*ALPHA* X (1) / (SQRT ( (P MAX**2) +(ALPHA*X (I) ) **2) ) -  R  B  91  using  a non-linear least  Bevington the  (1969).  respiration  were f i r s t Jassby  term  or n e g a t i v e i n t e r c e p t  Piatt  using  linear  (1976)., used  mathematical  i n the  PMX  v a l u e s r e p o r t e d a r e based hr-*  The  fits  data  were t h e  were  a l l  the  f i t of  bloom and  are  equal,  16, °C  was  net  advised  estimates f o r  by  these  PC1AX.  parameters.  productivities  evaluated  by  in The  ( ug C  with  P>.90 t h a t the  two  there  was  models,  ALPHA  c o n d i t i o n s a t any  indicated  that the  A t-test  18  (ug  conditions  20  o f the  light  although  difference the  u s i n g the  hypothesis. phytoplankton  fitted  standard  TANH  model  similar  between  temperature.  The  PHAX*s were l e s s d u r i n g  (for  parameter and  °C .... The  s a t u r a t i o n curve  little  given  a  given was  Furthermore, may (Piatt  have.an and  conditions  i n the i t  important 1976)  forcing  difficult  the c e l l  Jassby,  PMAX*s  significantly  post-bloom  5 made  the  temperature)  not  variability  (Sfi,TEMP,N0.3) d u r i n g E x p e r i m e n t this  a c t u a l and  were a l s o  the  In a l l cases,  o f the h y p o t h e s i s t h a t t h e  that t h i s  °C o r  calculating  search technique.  v a l u e s of  post-bloom  ,  confirm  the  The  indicated  self-shading on  on  (P>.10) between t h e b l c c m  conditions to  the  period.,  for  for  irradiance,  as  v a l u e s and  ,  e  models a r e summarized  PMAX, SIGHAA, were much l e s s  i n F i g u r e 96  different  resulting  output  a grid  excellent  post-bloom  post-bloom  f i t  using  7 cases.  bloom a n d  .R ,  e s t i m a t i o n cf the g r o s s  same.. F u r t h e r m o r e ,  d e v i a t i o n s of  data  squares  chi-square  the  in  at zero  in  ).  least  between  i n p u t and  outlined  f o r ALPHA and  regression  r e s u l t s f o r both  19,  reduced  including  The  Table  Chla)-*  procedure  parametric estimates  p a r a m e t e r s were t h e n The  f i t (LSF)  The  calculated  and  squares  size  and  effect as  well  92  as t h e temperature  and n u t r i e n t  status of the c e i l s .  S i m u l a t i o n o f the P h y t o p l a n k t o n  Dynamics  D e s c r i p t i o n o f t h e Model In using  order to t e s t the  simulations language The  the v a l i d i t y  experimentally  cf  the  determined  productivity  parameters,  a s e r i e s of  were r u n u s i n g SIMCQN, a s i m u l a t i o n c o n t r o l  V,  w i t h t h e model programmed  primary  phytoplankton  purpose  stock  time  command  i n FOBTB&N.  o f t h e s i m u l a t i o n s «as t o  over  model  from  predict  the primary  productivity  model a n d compare i t w i t h t h e a c t u a l e x p e r i m e n t a l r e s u l t s . operational of  the  (TEMP)  procedure  forcing and  of the s i m u l a t i o n allowed  conditions,  nitrate  concentration  to  phytoplankton  stock  productivity  (N03),  be  model  solar  read was  and  radiation  and in  This  'final  phytoplankton Iterations SIMCOM  and  stock  of was  this  1  was  for the  following  procedure  determined  then  by  was the  i Documentation f o r this procedure Computing C e n t r e f i l e iaBE:SIMCON.»  values  (SB), temperature  calculated derivative  value  The  phytoplankton  The i n c r e a s e i n t h e based  on  the  added t o t h e i n i t i a l  value i n order t o estimate the phytoplankton day.  the d a i l y  seed  initially.  then this  the  the  stock  used day.  specified number  as  after the  The  that  initial  number  of  interactively in of  days  i s a v a i l a b l e from  i n the  t h e UBC  93  experiment.  It  discussion, the  should  be  noted  s t o c k and a s s i m i l a t i o n  the. experimental  variables  CHLA  f o r a l l f o u r In  the  data averaged  the  situ  which EXTK  coefficient  i s  a  (EXTK)  function  photosynthetically  of  available  f o r t h e tank  (since  following  variables  for  r a t e , and c o r r e s p o n d t o  CHLA and ASS.  b a s i c s t r u c t u r e o f t h e model  extinction  average  in  PHYTO and P r e p r e s e n t t h e s i m u l a t i o n  phytoplankton  The  that  PBAV  represents the  stations. , included  using  PHYTO  the system  of  Biley's equation i n  (see  radiation  calculation  Chapter  was was  2).  calculated turbulently  The a s an  mixed)  using the equation: PABAV= with  0.5*Sfi/(EXTK*2) *  the.appropriate conversion  into  langley  estimated results  min-* .  using the  factor  following  GECM  eguations  ( t ~ * ) , was  determined  P  The  estimates  ALPHAI  were  However, b e c a u s e o f t h e l i m i t e d versus  TEMP •  parameters, productivity  curves,  ALPHAT at  from  the  Table  19.  = PMAX*TANH (AIPHAI*PAEAV/P13AX)  CCHLA i s t h e CABBQN:CHLA of  then  /CCHLA  PMAX = PTMAX*TANH(ALPHAT*TEMP/PTM and  the v a r i a b l e  experiments:  GBOWH = P  to transform  The g r o w t h r a t e ,  o f t h e *P v e r s u s I '  where  (1-EXP (-EXTK*Z) )  the  taken  ratio.„  directly  number o f d a t a estimates  (the i n i t i a l  slope)  AX)  of  and  from  p o i n t s i n the the  corresponding  PTMAX  (the  t h e o p t i m a l SB and TEHP f o r t h e g i v e n  c o n d i t i o n s ) , , were more d i f f i c u l t that  to predict.  literature  suggested  PMAX  temperatures  only marginally higher  A  approaches than  *P  survey a  maximum nutrient of  maximum  the at  20 °C f o r a S k e l e t o n e m a  94  costatum  population.  experiment  (Figure  seemed r e a s o n a b l e measurements possible. is  by  96,  first  of t h e  during  Table  values  system  phytoplankton conditions  as  relatively  (see  used  CCHLA  the  phytoplankton,  rate  i n t o the  the  system  phytoplankton the i n i t i a l (  0.30  for  to  experimental  - 1  a  data  in  from  10,0  PTMAX.  No  this  to  ca.  the  simulations  by  the  with  60  direct  v a l u e o f 45  was  of  a  nutrientT h e s e CCHLA  the  appropriate  results.,  a l s o used d u r i n g  phytoplankton  ca.  phytoplankton  1973a).  experimental  CCHLA  values of  in  during  15.0  (CCHLA) were  similar  Takahashi,  An  periods  of  concentrations.  the  growth  rate  of  the  c a l c u l a t e d as:  (GBOWB*PHYTO)-(FB*PHYTO) - (SINK*PHYTG)  of  the  equation was the  phytoplankton, as  w e l l as  continually b e n t h o s was  d e p l e t i o n of n i t r a t e .  day  dynamics,  of  t h e d e r i v a t i v e was  sinking  introduced  values  calculation  d(PHYTO)/dt =  to  in  steady-state  After  though  values  for  P a r s o n s and  indicated  intermediate  The  the  c a r b o n r c h l o r o p h y l l a parameter  bloom  were  periods  27),  approximations  community, i n c r e a s i n g t o depleted  e x t r a p o l a t i o n of  However, o t h e r e x p e r i m e n t s h a v e i n d i c a t e d t h a t  a f u n c t i o n of the  30  So  the  SINK  (  flushing  mixed,  some  t  -  rate. loss  apparent, p a r t i c u l a r l y Sinking  r a t e s of  0.10  were t e s t e d i n t h e s i m u l a t i o n r u n s b a s e d  estimates.  ),  1  was Even of  after day  on  - 1  some  95  Simulation The  Results  results  o f some o f t h e s i m u l a t i o n  are  illustrated  i n Figures  and  102 f o r EXP5A. data  rate  f o r s i x weeks.  1,0  day  f r o m EXP5B, t h e e x p e r i m e n t  ,  The s i n k i n g  the  bloom,  ( s i n c e t h e tank held The  constant  the  third  SINK  was v e r y  and  in  view  of  concentration the:  end  altered  this  0,0  day  f o r four  - 4  ;  days  and  then  t o 60 d u r i n g value stock  the  post-  o f 45 d u r i n g concentration  t h e f o u r t h week, CCHL was a l t e r e d  large  (Figure 74).  - 1  a s i n the  - 1  period),  r e s e t t o ah i n t e r m e d i a t e  the  at  set at  f o r t h e r e m a i n d e r o f the e x p e r i m e n t .  - 1  During  flushing  increase  in  the  inflow  CCHL was.then i n c r e a s e d  to  nitrate  t o 45  until  o f t h e 1,0 day-* FB p e r i o d and r e s e t t o 60 when FB was t o 0.5 d a y - *  .  derived  the  approximation estimated  had  clear during  i n Figure  timing  set  was a l t e r e d t o 0,3 d a y  As i l l u s t r a t e d those  t o 0,5 d a y  101  a p p l i e d to  t h e FB was  week when t h e a c t u a l p h y t o p l a n k t o n  was f a i r l y s t a b l e . 30  the s i m u l a t i o n s *  was s e t a t 3 0 , i n c r e a s e d  period,  Figures  t i e constant  r a t e was i n i t i a l l y  a t 0,1 d a y  CCHL r a t i o  bloom  During  with  and on Day 41, was d e c r e a s e d  experiment. after  97 t o 100 f o r EXP5E and  The s i m u l a t i o n a n a l y s i s was f i r s t  the:  - 1  runs f o r Experiment 5  from  of  by  the  the  97, u s i n g  'P  actual  simulation  versus  l a g during  PHAV, / T h e s e n s i t i v i t y parameters  in  0.4  after  day-*  I*  model  the f i r s t  a  initial  the  simulated  two weeks compared  was t e s t e d  bloom,  (PHAV)  the  was  The m a g n i t u d e and  by a l t e r i n g  t h e model., By i n c r e a s i n g SINK f r c m the  plus  reasonable  stock  (PHYTO)•  although  o f the system  parameters  data,  phytoplankton  o f t h e maxima were s i m i l a r  a one d a y t i m e  these  0.3 d a y  recovery  of  data with the - 1  to  the  96  phytoplankton  stock  by c a .  50%,  closer  t o the  98).  On  day  although  A  stock  out  predominant f o r c i n g oscillations indicated the  t h a t the  standing  variability apparent was  nitrate GBOWB often  variability  in  simulated  the  experimental  c o n c e n t r a t i o n and a  function  described  estimate  of  by  10.0  compared The  of  during  c o n d i t i o n s and  simulate  (Figure results  a c c u r a t e l y because of (0.1  day  magnitude  and  frequency  Simulation  runs using a constant  appropriate  4  to 0.5  flushing  of  N03  simulated  hr  - 4  damped of  N03  week,  not  system  a function of adjusted  so  the that  a s s i m i l a t i o n of The  on the  was  t h a t the  N03,  original  f o r the p r o d u c t i v i t y  of  EHYTO  experimental  100)  the  effect  stock  kinetics. 1  were  fourth  then  99)•  simulation  significant  uptake and  (ug C h l a ) -  lower FB*s  the  was  (Figure  or  indicating  i n the c a l c u l a t i o n  PTMAX=12. 0  in  phytoplankton  a c l o s e r p r e d i c t i o n of the with  oscillations  the  the  then  (Figure  _ 1  depletion  model c o u l d be the  c  ug  a  was  0.1  how  and  had  Michaelis-Menten  p a r a m e t e r PTMAX used provided  the  SB  reduced  SINK o f  approximated  bloom  growth  1  a  a c t u a l ; and  results  The  Chi  p l o t s was  i n SB  However,  simulation  a constant  which  initial  light-limited.  was  in  stock.  i n the  not  the  c o n d i t i o n s . , The the  ug  week was  pcst-blccm  not a s c l o s e l y  periods  after  third  maintaining  feature of  the  o f 54.2  bloom, the  are  significant  model p o i n t s  value  hand, by  the i n i t i a l  phytoplankton  middle of the  t h e PHYTO(max) f o r t h e  experimental  the other  after  - 1  i n the  or higher  (Figure standing  values  of  97) stock  PTMAX. ,  c f EXP5A were more d i f f i c u l t  to  the  of  day-* the  changes )  which  parameters  value  r a t e s and  i n FB  of  10.0  reasonable  and  would in for  the  use  alter the  system.  PTMAX,  estimates  the  with of  the  97  slaking  rates  0.5  -  day *  f o r Period  about t w i c e the 5.0  3),  nutrient concentration ug G (ug C h l a )  Chla)-*  (ug C h l a )  Period  t o CCHL d u r i n g  period.,.  As  3.  phytoplankton  post-blccm  decrease  sensitive  to  in  stock  Period  levels  of  a  Figure  concentration  i n SB compared  I f SINK was  as i l l u s t r a t e d  grazing  pressure.,  •crashed* to  effect when  phytoplankton  was o b t a i n e d a  with t h e h i g h  the s i m u l a t i o n  large  tank  steck  reduction  run.  ,  o f the  reached  in  the  »as flushed  inflow  103, t h e r e even  the  negative  extinction Hhen t h e  primary  This scenario  was a  with  the  the  system  corresponded a  similar  Parsons  (1972)  a t a r a t e o f 2.0 day-*  and t h e  l o s s r a t e o f 2.0 d a y *  experimentally  less  i n t o the  NC3 c o n c e n t r a t i o n .  was i n c r e a s e d t o 1.0 day-*  an o v e r a l l  was  stock  T h i s r e s u l t e d from  of t h e consequent  during  closely  of N03.  i n Figure  additional  rate  more  system  t h e phytoplankton  of the phytoplankton  grazing  increased  50 ug C h i a 1~* i n r e s p o n s e t o t h e h i g h  coupled  was  resulting  to the combination  recovery  coefficient,  60  (GBA2E) o f 0.5 day-* was i n t r o d u c e d  o f N03.  effect  of  the  and magnitude  significant  feedback  value  101,  (PHAV).  pattern  on Day 27 a f t e r  ca.  to  2 and t o 5.0 ug C (ug  c h a n g e s i n FB and low i n f l o w c o n c e n t r a t i o n s  model s t a r t i n g  with  bloom a n d was r e s e t  As s i g h t be e x p e c t e d , t h e  a g r a z i n g term  was  (PH¥10) d i d n o t show a s i g n i f i c a n t  and r e c o v e r y  changes  decreases  2;  Period. 2 and CCH1=75 f o r t h e 0. 10 day-*  t o 0.6 day-* •, t h e s i m u l a t i o n a p p r o x i m a t e d PHAV.  the i n i t i a l  Similarly  illustrated  simulated  PTKAX  which  o f t h e s y s t e m , PTMAX was d e c r e a s e d  hr~* during  - 1  f o r Period  - 1  r e s u l t e d i n a PHYTO(max)  hr~* after  - 1  hr~* during  assigned  1; 0. 1 d a y  a s l a r g e as P H A V ( m a x ) S i n c e  t o 8.0 ug C  FB  (0.25 day-* f o r P e r i o d  -  by Brown and  .  98  phytoplaokton  stock  was  reduced t o z e r o .  9 9  CHAPTER 9.  The  COMPARATIVE DISCUSSION OF  experimental  importance  of the  determining various  the  results of  flushing  flushing  rates  of  proportional  changes  in  and  of  phytoplankton  productivity  at a l l the experimental  1.00  day-*  )  was  s o u r c e o f seawater experiments, autumn  (av N03  The a  determined stage  ca.  = 2k  uM  upwelling  with  restrictions  on  an  source.  artificial primary  experimental  tanks  particularly  in  sinking  rate  constant this  the  r a t e o f the  The was  a  forcing  the  primary  (0.25  day  - 1  nutrient-rich _  during  t  conducted  rate  community  the  first  the  i n the  six-week weeks  phytoplankton,  r a t e , c o n t r i b u t e d to the  with  and  over  of  water in  due  and  column  the  the  oneUnlike  to  dynamics  .  significant Hithin  the by of  the the d u p l i c a t e  pre-grazing  relative  time.  )  - 4  which c o u l d be a c h i e v e d  between  three  day  light-limited  reproducibility excellent  (0.5  c o n d i t i o n s (EXPI).  was  intensity  during  variability  was  l  in  The  rates  N  inflow  available  changes  stratified  primary  upwelling depth  which  in  system  the l i g h t  system  1  uM  The  subseguent  the  u s i n g a deep 20  in  ).  4  the dynamics of the  culture  w e l l as  between t h e f l u s h i n g  the other experiments,  the  N 1~  with  s t o c k and  flushing  which a v e r a g e d Experiment  i n the  communities.  by  interaction  constant  as  enhanced  except  the  chains.,  difference  phytoplankton  composition  to  food  temperature,  productivity,  indicated  turbulence of the system  a  r a d i a t i o n f o r primary the  have  herbivorous  provided  c o n c e n t r a t i o n s of n u t r i e n t s  EXPERIMENTAL RESULTS  t h i s study  r a t e and  dynamics  THE  The to  period,  differential the  constant  i n c r e a s e i n CHLA' each  depth,  the  100  phytoplankton and  stock  reached  a 30  stratification  of the  response  of  ug  the  were d o m i n a n t a t t h e The  complicated Navicula  by a  into  i n Tank  of  solar  and  {such  and  PAB  of  the  culture  a l a r g e degree  may  have  with  of  4  introducing  phytoplankton with  an i n i t i a l  experiments  almost  at  bottom  community a  were  filamentous  t a n k s . ... T h i s  light  that  The  diatom  by  oxygen  intensity  at  in  the  N03  occurred  both  depth,  low  high  micro-nutrient the  flushing  on  Day  the  The  Experiments Despite  5 i n a l l the  reach a  develop,  experiments,  the  of  mixing,  radiation.  turbulence.  ; in contrast,  day-* d i d n o t  limiting rate  to  eliminated during  between  most o f  of a r t i f i c i a l  solar  non-  forcing conditions  seme  lack  artificial  FB=0.25 day-* 1.0  a t the  stratification  of  totally  TEMP and  bloom  the  periods  was  i n SB,  in  r e p l a c e d n i t r a t e as  thermal  stratification  variability  depth  of  were n u t r i e n t - l i m i t e d  an i n d i c a t i o n  combined  by  the  with n a t u r a l  - 1  5  the  of  experiments  0.25  and  in  change  Micro-flagellates  growth  the  i n some o f t h e e x p e r i m e n t s .  during  a  mats, , s u p p o r t e d  temperature  t h e r e was  particularly  The  with diatoms  cf  nutrient  allowed  water column.  phytoplankton  reduced  t u r b u l e n t systems  ,  the  averaging  B.  a s v i t a m i n B12)  day  month,  TEMP..  the  oscillations  as a f u n c t i o n  and  surface  further  da aped  promoted  flocculent  r a d i a t i o n and  thetime,  also  in  the  c o n t r a s t , the  turbulent  for  l  effect,  large  particularly  l~  s u r f a c e compared  at  and  a  of  one  p r i m a r y community  wall  •vacuoles*,  after  system  dynamics  species  developed  In  Chi  to the decrease  station..  a series  steady-state  approximately  composition  displayed  the the  experiments  CHIA  naximum u n t i l  in Day  the 8.  101  In a l l cases, situ  nitrate  function  the bloom  concentrations.  of  the  concentration) of  this,  was.coincident  flushing  4  and  culture  and  the  decrease  productivity variability probably grazing  correlation In  o f t h e u p t a k e o f oxygen  these  the  h e r b i v o r e s , the c o r r e l a t i o n by  experiments  the  was  herbivores primary  i n m a g n i t u d e and  (Experiments  a r e s u l t o f the lower f l u s h i n g  when t h e f l u s h i n g  , provided  that  4 indicated  2  and  3),  r a t e as w e l l as t h e  that  r a t e was d o u b l e d  the  system  the l o s s o f phytoplankton 0.5  rates greater  than  composed  diatoms.  of  provided  supporting  composition flushing  increased A  grazing  stock.  (ASS) were a l s o much g r e a t e r  during  spite  for  the  same  c o n d i t i o n s o f SR, TEMP and NC3, t h e p h y t o p l a n k t o n  minimize  -i)  and phytoplankton  r e s u l t s i n Experiment  forcing  day-*  no  was a h i g h  N03  pressure.  The  doubled  with  a  in all  1  i n CELA by g r a z i n g . , The s t a n d a r d i z e d  rates  as  associated  c f SR, b u t i n  cultures  5), there  with i n s i t u  as a result  the  was  30 t o 40 ug C h i a 1 ~  In the primary  (Experiments  one-stage  (and  and m a g n i t u d e  between t h e oxygen p r o d u c t i o n  reduced  rate  g e n e r a l l y r a n g e d between  pressure  the d e p l e t i o n o f i n -  T h e m a g n i t u d e o f t h e bloom  and t h e t i m i n g  of t h e experiments.  with  day  - 1  The  evidence  of the primary  component  periods  0.5 d a y  tcrbulently  - 1  t o 1.0  mixed  to  t o t h e b e n t h o s . . At f l u s h i n g  , t h e primary pigment  community  was  r a t i o s and C o u l t e r  that there  was l i t t l e  community a t c o n s t a n t ,  r a t e s , although  during  was  frcn  stock  the proportion  of  mainly counts  change i n the high  (>0.5 day  nano-flagellates  o f low i n f l o w N03.  a n a l y s i s of gross  primary  p r o d u c t i v i t y during  E x p e r i m e n t s 4 and 5 i n d i c a t e d t h a t a t f l u s h i n g  r a t e s o f 0.5  day  102  -  1  or  greater, a large proportion  productivity flushing  was  rates  fraction  in  *lost* (0.1  the  day  one-half  probably  a  as  assimilation 35%  and  - 1  fraction  of t h e  was  estimates  of  parameters  function  of  the temperature  system  indicated  productivity the  two  had  a  two  become  would  H03  like  nitrate;  for  controlled  *P  i n SR  in the  versus  f i t  simulation  of  were  magnitude  TANH  model w i t h  (the e x p e r i m e n t ALPHAC  a  the  TEMP,  of  experimental  based). the time  were  The  the  estimates  model  phytoplankton  a  used  p e r i o d s when t h e  of  aid  direct as  three  between  significant.  data  at  system, did  various in  not  provide  However, t h e  of  EXP5B  PTMAX dynamics  illustrated was  least  phytoplankton  PMAX,  productivity  the  combined  non-linear  the  the  standardized  difference  in a  a  days  Although  i n the case  predicted stock  to  nutrient  variability  success, p a r t i c u l a r l y  f o r which  and  f a c t o r a l experiments.  model,  20%  system,  the  I*  e s t i m a t e s o f : t h e parameters p r e d i c t e d from sguares  from  statistically  and  rates,  to obtain  t c the experimental  the n a t u r a l v a r i a b i l i t y  conditions  ranged  system  difference  not  15%,  t e m p e r a t u r e s . , The  nitrate-limited.;  however,  concentration  flushing  culture  the  lower  respiration  c o n d i t i o n s of the  of  at  .  - 1  r a t e s between t e m p e r a t u r e s ,  one  30%  primary  r a t e s was  attempted  given  and  a significant  concentrations of  with  for  and  the  The  lower  day  experiments  n u t r i e n t c o n d i t i o n s was  Ideally  inflow  I»  with  ).  in situ  0.5  before n i t r a t e - d e p l e t i o n  the  results  lower  of  flushing  more v a r i a b l e  versus  after  dayr* high  r a t e s of a t l e a s t  t o 60%)  compared  value f o r the  VP  status -  0.25  with  the  result  at flushing The  by e x u d a t i o n ,  systems  approximately  < 5 0%  and and the  independent  103  o f S H . _ The b e s t f i t f o r obtained  using  determined  a  from  the  constant  the * P  in situ  Although  t h e p a r a m e t e r CCHL  used  rate  -  * ), only  with  post-bloom  steady-state  phytoplankton  stock. ,  factor  nutrient-limited. turbulent  the  Consequently, flushing  to  to  the  model  30  for  during  the  close prediction  o f the  sinking  rate  was  r a t e and  was  in  a  the  non-  significant in  the  phytoplankton.  f o r EXP5A, w i t h  also  became  more  of  accurately.,  a  the v a r i a b l e  and m a g n i t u d e o f  Ecwever, function  magnitude and v a r i a b i l i t y  the of  were more  other  F B r and difficult  r e s u l t s o f t h e h e r b i v o r e growth and s u r v i v a l d u r i n g t h e  experiments  indicated  scallops  was  two main f e a t u r e s .  enhanced i n b o t h  systems with high f l u s h i n g the  were were  were  the o s c i l l a t i o n s  sere  the  predict. The  the  tank.  estimates  the  heterogeneity  as  the  determining  60  was  d e p l e t i o n , and 45 d u r i n g  flushing  simulation runs  stock  estimates of t h e i r  in  measurements:  a  SINK  was  with  f o r the primary  parametric  systems,  cultures,  spatial  phytoplankton  agreed  sufficiency,  nutrient  At t h e l o w e r  the  in  of direct  provide  r a t e , d i d not p r e d i c t  parameters  and  f o r a b o u t f o u r days a f t e r t h e s y s t e m  one-stage  determining  EXP5B  three r e p r e s e n t a t i v e values  These  Even i n t h e t u r b u l e n t  for  f o r PTMAX, which  important  nutrient  period..  sensitive  the  was  p e r i o d with  sufficiently  significant  data  v a l u e s o f ASS c a l c u l a t e d  conditions  initial  tuns  o f 10.Q  I*  i n the s i m u l a t i o n i n l i e u  bloom  a  ( t  value  versus  maximum  growth  simulation  herbivores  were  rates  First,  the  t u r b u l e n t and  (> 0.5 day-*-  l o c a t e d a t a depth  growth  of  non-turbulent provided  o f 1.0 m t o a v o i d  that high  104  light  intensities.  that  the  {av  g r o w t h was  TEHP=9.5 °C  maximum  of  (FB=0.75 during  EXP2B  temperature  stock  to  alleviating  the  However, appeared  with  IV) , their  the  density,  provided  as  the  i n the  Crassostrea determined Crassostrea day-*  .,  low  during  were low  the  growth 2.0  source  density  gigas  the  However, i n t h e  m  a  EXP3B  was  poor  increased  and  decreased  growth  1. 18  g/zoo/week  cm.,  excreted  other  ammonia  cultures  I  of  phytoplankton  determined highest ca.,  by  the  rates  were  20.0  1/day  potential filtration  feeding  experiments  D u n s t a n , 1973b)»  experiments  to  a function  of  received  filtration  in  productivity.  (Experiments  was  and  was  stock.  type  which  of  It  one-stage  oysters  The  of  the  priitary  about 6.5'1/hr i n a oyster  during  1.0  5.0  the  the  Tenore  reached  of the  fcr  EXPIV., T h e s e  with  EXP1  a flushing rate  to  of  (which  tank,  average  was  at  r a t e of  cm  tank).  E X P I I and  1972;  °C  C r a s s o s t r e a .gigas was and  during  system.  phytoplankton  primary  (Balne,  14.5  at depth  during  size,  compared  that  of  s i g n i f i c a n c e of  of  food  apparent  rates  the  by  their  r a t e of the  oyster  from  growth  rate  flushing  per  culture  indicated  scallops  a result S3  t w o - s t a g e c u l t u r e of  growth  temperature  the  as  nitrogen-limitation oyster  the  )  a maximum  i n size  to be l i m i t e d  During  EXP3B  c u l t u r e systems with  estimate  the  and  a s u i t a b l e environment  oysters ranging  impossible  by  temperatures  ), increased  the  gigas  limited  day-*  i n this  day-* p r o v i d e d  EXP1  growth e n v i r o n m e n t even  day-*  contrast,  Crassostrea for  in situ The  (15.6  of  growth r a t e of  (FB=0.25  phytoplankton  0.25  at  )•  -  probably  ) . , The  16.8%  day *  In  a comparison  rate  for  system with in this  5.5  for flalne cm  a FB=5. 0  study  (EXPI  105  to by  EXPIV), the  this  flow  r a t e from  feeding  primary  tank  to l e v e l s  of  system..  In  in  the f l u s h i n g r a t e of  rate would 2.0  in  day  a  be  increased,  class,  i n weight  by  c_a.  four  major f e a t u r e s . ,  the  significant  ( 8 . 1 % - p e r week). day-*  with  flushing system  herbivore  r a t e of 0.10 provided  the  but  day-*  a  of p h y t o p l a n k t o n  herbivore  F u r t h e r m o r e , the  tanks  high  tanks  significant  f a c t o r i n the  increase  primary  when  21%  i n one  herbivore First,  week. ,  experiments  the  least  growth  during  rate  w h i c h was  food  rate of  growth  This source,  experiment  reduction  of  the  of  (>23  1.0  promoted by  the  °C  )  was  a  primary but  maximum a t t a i n a b l e f l o w  this  of  during Experiment I I I  qualitative  the  FK  i n Experiment I I ,  tank.  and  in  found  primary  limited  during  the  the  r a t e s were  r e s u l t a n t temperature  herbivore  limited  increase  t a n k s were f e d a t a  in the  suitable  an  growth  a n a n o - f l a g e l l a t e community  concentration of  The  to  'wash-out* o f  particularly  between t h e  was  order  since  highest  EXP5 i n d i c a t e d two oysters  In  t a n k s was  t w o - s t a g e c u l t u r e , t h e volume o f  terms of s i z e , the  comparison  herbivore  tank.  only causes a  - 1  oysters increased &  primary  have t o  the smallest s i z e  the  the  the  the rate  oysters. in  probably  meat t o s h e l l  the a  weight  ratio. , Secondly, averaged  the  greater than  the  phytoplankton  that  e i t h e r a 0.5  primary The the  o y s t e r growth i n the  tank  highest  10%/week.  community day-* or  provided  an  was 1.00  culture  experiments  these  composed  diatoms,  day * -  e x c e l l e n t food  during  three  In e a c h o f  average growth r a t e s of  two-stage  other  of  flushing source  experiments, indicating  rate  f o r the  of  oysters.  13.5%/week were a t t a i n e d  Experiment  II.  the  During  in this  106  experiment, the Crassostrea  qiqas  2.0  B  day'  the  frcm  highest  costatum to  .  the  oxygen  growth  t o be l i m i t e d  by  indicated  that  g r o w t h was  not  An  temperature  in situ  25  growth r a t e  the  decreased oysters  after  3.5  another  than  Ladysmith  Harbour,  locations  tested  would  field the  density  oyster  *as  were h i g h  i n the continuous  four  tanks  d i a m e t e r X 1m  (6m  same p r o d u c t i o n  a s one  feasibility  system should  be  of  assessed  of  tank  appeared The  required  data  so  At be  indicated  that these  achieved. that  the  that  the  o f c a . , 60  g  Experiment I I c u l t u r e system.  measured  were by  the  Quayle of  oysters  carefully.  (1971)  the  r a t e s of  c u l t u r e system,  Quayle*s  greater  or f o u r years  growth  would  15%  be  at  were the  Crassostrea approximately  required to  rafts.  at  sixteen  Marketable oysters  of three  deep)  growing  qiqas  respiration.  a marketable s i z e  Columbia.  However, a l t h o u g h  according  concentration.  o y s t e r growing area  years, instead  for  17.71/week c o u l d  the  rates  best  which  i n c o r p o r a t i n g the f a c t  be  in British  a f t e r two  sites.  economic  high  Skeletonema  of c r a s s o s t r e a  increasing size,  gigas  the  growth  ,  growth r a t e s i n t h i s c u l t u r e s y s t e m maximum  other  results,  with  °C  by  of  contained  concentration.  food  r a t e s of  months u s i n g  the  obtained  the  growth  which  were s u f f i c i e n t  CHLA/day/g by  ),  dominated 20.1  f o r the  levels  ug  experimental  The  averaged  phytoplankton  limited  extrapolation of  CHLA  of o y s t e r s i n the  the  levels,  (FB=1.Q d a y - * of  (1969) i s o p t i m a l  However, the  feeding  tank  concentration The  Quayle  , and  primary  Mere f e d a t a f l u s h i n g r a t e  achieve  Therefore  the  in a controlled culture  107  CHAPTER  10.  SOMH&BY JJJC  COHCLDSJONS  Five continuous c u l t u r e experiments the  hypothesis  enhanced optimal  that  by u s i n g  optimal system  limited,  and  radiation  with  structure  compensation  system  0.33/day  phytoplankton  a maximum  was c a .  of  the  solar  stock  of  water c o l u m n a t  c f 25 uM H "1-*  .  Based on a  compensation  the f l u s h i n g r a t e  could  depth at this  be  reduced  to maintain  t h e same  concentration.  diatom the  Skeletonema  growth  rates  of  with t h e outflow from  temperature °C  was o p t i m a l )  costatum  1.0/day f l u s h i n g r a t e  2. Q/day  (20. 1  to  nitrate-  3.0 m e t r e s i n d i c a t i n g t h a t  when t h e h e r b i v o r e  gigas  was  phytoplankton  i n t h e one metre  achieved  The  be  that:  conditions  i n a 3.0 metre impoundment  at  maximum  could  of the ecosystem  system  forcing  g r a z e d when f e d t o t h e c o m m e r c i a l The  The  i n t e n s i t y o f 18*  nitrate,  chain  ecosystem  natural  concentrations  light  of i n f l o w  2m.,. T h e  chains  test  t o maximize p r o d u c t i v i t y i n a  1.0/day.  was a t t a i n e d  _ 1  nitrate  the  was  and t e m p e r a t u r e ,  ug C h i a 1  inflow  to  and s p a t i a l  flushing rate  turbulent  level  food  p r o d u c t i v i t y . , The r e s u l t s i n d i c a t e d  1. . T h e  for;  i n bivalve  to  a deep n u t r i e n t - r i c h s o u r c e o f s e a w a t e r and a n  flushing rate  maximize  60  production  were c o n d u c t e d  and  r e s p i r a t i o n requirements.  the  tanks  and was  juvenile  for  the  growth  levels  gigas  oysters  were f l u s h e d  at a  of  the  selectively  Crassostrea  the p r i m a r y tank  oxygen y  oyster  dominated  rate  .  were of  (FE= 1.0/day) . Crassostrea  were s u f f i c i e n t f o r  The growth r a t e o f t h e o y s t e r s  was  108  a  f u n c t i o n of t h e i r  the ug  size,  maximum p e r c e n t  with  increase  the smallest  size  group  having  i n w e i g h t , and an a v e r a g e o f  C h i a d y * p e r gram o y s t e r  was r e g u i r e d  -  to achieve  25  maximum  g r o w t h r a t e s o f 18%/week.  3.  In  terms  mariculture, culture  of the  system  the growth  were  rates  ca..  •optimal*  field  this  (7500 l i t r e s )  size  application in  size,  and  first  growing s e a s o n  could  provide  the  152 g r e a t e r  location in British  gram  o f the r e s u l t s t o o y s t e r optimal  than t h e r a t e s i n an  Cclumiia. ,  support  continuous  A  system  250 c y s t e r s / m  a marketable crop  (when t h e o y s t e r s  of  o f a 10  3  a t t h e end o f t h e  are  ca. ,  1.5  years  old) .i  4. . A t in to  lower f l u s h i n g rates  the standing the  stock  decrease  flushing lower  rate,  of  Chaetoceros sp.  5.  flushing  in  rate.,,  temperatere,  sere  stock  the  levels  community  from  and  although the i n s i t u favourable  at  was l i m i t e d a  change  this  by t h e  in  the  .  chain,  provided  that  were grown a t a d e p t h o f 1.0 light  proportion  Skeletonema costatum t o  H i g h e r f l u s h i n g r a t e s a l s o enhanced  s c a l l o p food  was a d e c r e a s e  direct  t h e growth o f t h e o y s t e r s  phytoplankton  composition  of phytoplankton  in  c o n d i t i o n s , such as  (0.25/day), there  intensities.  Maximum  w e i g h t p e r month were a t t a i n e d  the p r o d u c t i o n  t h e Chlamys h a s t a t a  metre growth  to  avoid  high  o f the hericia surface  r a t e s o f 16.8% i n t o t a l  a t a f l u s h i n g r a t e c f 0.75/day  109  when  the  in situ  phytoplankton  temperature  concentration  was  {  29  14.5 ug  limiting  growth.  However, under t h e s e  growing  species  such  preferable  €.  at  f o r marieulture  very  (2.0/day), and  high  other  rates stock  such  The p r o d u c t i v i t y  was  stratified  was washed  as  Navicula  of Navicula  and  a significant  pressure  of the i n s i t u  of  a  useful experimental  natural  phytoplankton  incorporating productivity predicted the with  and measured reasonable  phytoplankton  c u t o f the t a n k dominated the  levels;  in  two  when t h e  proportion o f the  and s e c o n d l y ,  continuous  tool  when the  for  accuracy  system  f o r examining t h e dynamics  determined  values  culture  a  simulation  model,  parameters f o r primary  the  forcing  conditions,  t h e p a t t e r n and magnitude o f  s t o c k , and c o u l d be used t o examine  various flushing  system  h e r b i v o r e s reduced  communities.  experimentally  with  would be  t o low l e v e l s .  In c o n c l u s i o n , t h e l a r g e - s c a l e provided  faster  primary  ,  light  the  stock  a  also increased  s t o c k sank t o t h e b e n t h o s ,  phytoplankton  the  ) was n o t  1  yessoensis  the  phytoplankton grazing  -  conditions,  of  s i t u a t i o n s in.response, to higher  system  a l  and  production.  flushing  species,  Chi  Patinopecten  the phytoplankton  attached  system.  as  °C  ecosystems  r a t e s , s i n k i n g r a t e s and g r a z i n g r a t e s .  110 Table  1.  S T O D Y  Relevant c u l t u r e s t u d i e s o f marine o r g a n i c p r o d u c t i o n i n terms o f t h e i r e x p e r i m e n t a l d e s i g n and c o n t r o l (*) jSOLE 01 |0EERATICK  TEOFRIC SCOPE  jOPERAl. | TIME  CCNIBCL |SPACE  COMMENTS  3  Field 1 19731 Gross e t a l . , J F i e l d 195G| Field I . I a T. E» , ] 1970 j LSC Takaiashi | et al.,1975| Baab e t a l , J LSC 19731 McAllister | LSC et al.,1961| A n t i a e t a l . , 1 LSC 1963J Brown and J LSC P a r s o n s , 1972| LSC Goldman | e t a l . , 1S75J Malcne e t a l . , | LSC 19751 Strickland | LSC e t a l . , 1969i LSC Ansell e t a l . , J 19631 T e n o r e e t a l . , j MSC 1973J MSC Dunstan and 1 T e n o r e , 1972J Dunstan and ] MSC T e n o r e , 1S74| SSC T e n o r e and j D u n s t a n , 1973al E p i f a n c and j SSC Mootz, 1976 j H a l k e r and J SSC Zahradnik,1976] SSC Kirby-Smith S \ B a r b e r , 1S74j SSC Ketchum | e t a l . , 1949| Haynes,  j 1 | | l J | | 1 J 1 J f  |  |  |  | | J J  i i \  i  1 J I  |  1 I 1 I 1 | | 1  N a t u r a l 1 none | A. T. | L a k e s y s t e m f o o d webj Scottish loch Natural t batch | f o o d webf inorg 1 | 750 a c . t i d a l cys 1 semi-c impound ment Salmon n a t u r a l J I Natural 1 batch J 2.5X10m p l a s t i c f o o d webj i n o r g | cylinder P. Comm 1 c o n t ] Two-stage C u l t . 45,0001 P.tank Oys inorg | 1 P. Comm J none . ] isc ne | P l a s t i c s p h e r e (121 cu m) 1 | A.I. J P l a s t i c sphere P. Comm 1 none (12 1 cu m) 1 j OPVI . 1 1800 1 tank P. , , C C M 1 c o n t ] 1 inorg P. Comm 1 c o n t j A. T. J 2000 1 tank inorg ] 1 J A.T. | 2000 1 tank P. Pop 1 cont 4 day e x p s . | 1 inorg P. Pop | A.T. J 3m x 10m tank 1 batch inorg | I tank p . pop 1 b a t c h J A. T. j 1000 1 1 1 inorg | 760 1 tank Cys, 1 cont 17 - 23 Deg C. Sea 11 ! p. CCBffl J P. 'Comm 1 s e m i - c j A.T. | 400 1 t a n k 2 0 - 2 6 Deg C. , org | \ 400 1 tank P. Ccmro I s e m i - c . | org 1 ! ] 9 1 trays Oys, 1 cont Scall 1 p. comm j | 100 1 t a n k oys 1 cont (xecircilat.) i p. pops | 10 1 raceway Oys cont i I (. Ix. 1x1. 0 m) 1 nat.comm] S ca 11 3 1 racevay j 1 cent (. 3X.2X.C5 m) 1 nat.comm| P. .Pops 1 s e m i - c J A.T. | 8 1 f l a s k inorg j 1  * SCALE CF OPERATION: LSC ( l a r g e s c a l e c u l t u r e - >1000 1) ;MSC (medium s c a l e c u l t u r e - 100-1000 1 ) ; SSC ( s m a l l s c a l e c u l t u r e - <1C0 1) TEOPHIC SCOPE: P. ( p r i m a r y ) ; Pop ( p o p u l a t i o n ) ; Coram(cemmunity); Oys (Oyster); Scall(Scallops) TIME (Bate o f a d d i t i o n ) : b a t c h ; s e m i - c ( s e m i - c o n t i n u o u s ) ; c o n t ( c o n t i n ) TIME (Form o f n u t r i e n t ) : i n o r g ( i n o r g a n i c ) ; o r g ( o r g a n i c ) ; P. ( p h y t o p l . ) , SPACE: A . T . ( a r t i f i c i a l t u r b u l e n c e ) ;UPW(upwelling) ; b l a n k (net s p e c . )  111  Table  2.  D e s c r i p t i v e s t a t i s t i c s summary o f s e l e c t e d v a r i a b l e s f o r EXP2A, i n c l u d i n g a breakdown i n t o t h e p i e - g r a z i n g and g r a z i n g p e r i o d . Day 37 and 40 .measurements f o r CHLA, PBGC and ASS have been e x c l u d e d f r c m t h e s t a t i s t i c s f o r a d i r e c t c o m p a r i s o n w i t h EXP2E. The s t a t i s t i c s f o r CHLA and ASS a r e a l s o g i v e n f o r the g r a z i n g p e r i o d c o r r e s p o n d i n g t o EXP3A. PRE-GRAZING (t=1,6)  TOTAL EXP1EIMINT (t =1,41) V A fi  1 STN  SR SAL TEMI  J  NC3  NH3  I  1 I  s  | 1 1 | 1 1  M E '0 I  I  E 0 I  1 1 1 1 1 I  OXY  SAT  CBLA  1 1 ] 1 I 1 j 1 i 1  ASS  I  | I  |  MEAN  S. D. |  6  380.  165.  6 6 6 6 6 6 6 6 6 6  9.6 0.50 3.47 14.4 13.3 2.79 13.2 2.66 3.88 14.7 1.72 23.3 15.4 10.ce 15.0 10.49 S.78 15.4 15.0 10.04  .35 | 35 35 j 35 j 35 l 35 35 35 | 35 35  6 6 6 6 6  7.09 9.37 9.22 9.08 9.72  0.17 3.57 4.01 3.S5 3.29  6 6 J 6 .| 6  8.8 13.2 14.9 8.5  10. 12 17.60 19.71 10.06  3 3 3  3.5 3.3  2.27 1.67 1.27  MEAN  S.D. |  41 41 41 41 41 41 41 41  410. 28.3 11.0 17.0 16.2 15.4 17.0 18.2  114. 0.61 G.87 2.60 2.26 1.62 2.61 3.62  J  | | I J  | j |  H  s  K  B  o I  s  M E 0 I S B  B  s B  1 1 1  E o  1 1 1 | 1 3 1  I  N  N  s  i  J EBCD  I I 1  1  s  M E o  s  M E S H  i I  B  | 1  M E  s  | 41 41 | 41 | 41 | 41 I 41 I 41 I 41 | 41 | 41 I 41 I 11 | 41 I 41 I 18 J 18 I 18 I 18  I  | \ | \ I  14 14 14 14 14 14  0. 77 0.43 0.50 0.42 0. 42 0.3 2 0. 64 0.48 0. 47 0.31 7. 50 0,39 11. 41 2.60 11.76 2.82 2.76 11. 72 11.26 2.24 82. 5. 1 141. 33.2 113. : e C 140. 34.1 6.4 2 5.2 7.2 10.83 9.5 11.88 6.50 5.6  20.2 28.6 32. 1 4.9 5.2 4.1  GRAZING PEE.IGD (t=7,41) N  MEAN  S.D.  410.  112.  11.2 17.5 16.7 15.8 17.4 17.3 G.5 0.4 0.4 0.3  G.68 2.17 1.76 1.02 2. 17 3.C8 C.59 0.34 0.55 C.36  35 35 I 35 35 35  7. 57 11.76 12. 19 12. 17 11.52  C .06 2.28 2.38 2.28 1.S6  12 12 | 12 | 12 I 7 I 7 I 7 7  . 3.3 4.2 6.8 4.1 4.3 5.4 6.2 5. 1  2.48 3.35 4.21 3.46 2.55 3.81 3.S4 3.86  5.3 5.7 4.5 5.7 5.8 5.0  2.76 1.91 2.91  35  | | | | | I  | | | J  | | i  j | f  2.3 5 | 1.8 8 i 2.15 |  2.7  I  11 |. 11 I 11 I 6 I 6 I 6 .  112  3.  Table  D e s c r i p t i v e s t a t i s t i c s summary o f s e l e c t e d v a r i a b l e s f o r E X P 2 B , i n c l u d i n g a breakdown i n t o t h e p r e - g r a z i n g and g r a z i n g p e r i o d . See Appendix 1 f o r a d e f i n i t i o n o f t h e v a r i a b l e names. The s t a t i s t i c s f o r C H L A and ASS a r e given f o r the corresponding time p e r i o d s i n E X P 2 A .  TOTAL EXPEEIKENT (t=1,34) VAB  |  STN  SB SAL TEMP  |  I  I  I  i  tj I  NC3  OXY  J J 1 J  B G I  J  s  1  | |  i i 1  1  |  1 1 SAT  CflLA  I  1  PBOD  ASS  i  1 1 1  i  1 ASS  |  N  34 34 34 34 34 34 34 34  120. 28.3 10.8 17. 2 16.4 15. 1 17. 0 18.8  1 4 7 . .] 0.67 | 0.86 ] 2.77 | 2.58 | 1.68 j 2.72 | 3. 58  10 10 10 10 10 10 10 10  M  |  E  1 | |  34 34 34  6.  I | | | | | | | |  34 34 34 34 34 34 34 31 18  | |  18 18  I  18  oI s M B  o I  M E  1  l  S.C.  G.71 0.58 0.67  I 1  1 1  I 1 | 1  M E AN  34 34 34  s  1 I  1 | |  N  1  1  i]  CHLA  s M  E 0 I c M  1  NH3  i  1  s M  cB s  PHE-GBAZING (t=1,10)  50 0.65 7.52 11.45 11. 57 10.51 11.39 81. 141. 140. 12 5 . 6.8 9.7 10.3 8.8  0.42 0.49 0.44 0.31 0.61 0.52  | j | | |  2.79 2.94 3 . 10 2.43 6.5 34.6 35.9 37.2  J  9.69 10.95 10.83 9.82  j |  | ] |  9 9 9  I  9 6 6 6 6  E G  M E  s  M  E  \  s  J i  M B  | | |  14 14 14  1 I f  14 14 14  MEAN  S . D.  |  390.  172.  i  9.9 15.4 14.3 14. 0  0.61 3.11 2.59 2.30 3 . 19 2.OS  I  1-5.3 22.6  7.19 12.03 12.11 12.07 11.69  N'  | j |  0.29 4.29 4.31 4.33 3.72  MEAN  S.  24  4 20.  13 9 .  24 24 24 24 24 24 24 24 24 24  1 1.2 17.9 17.2 15.6 17.7 17.2 0.5 0.4  0.64 2.29 2.09 1.11 2.20 2.78 0.45 0.40 1.54 0.43  24 24 24 24 24  9.0 13.7 14.0 12. 1 10.6 11.7 12.0  11.1  13.00 14.04 13.65 12. 45 16.05 16.06 15.72 15. 33  |  2.39  3 . 13 -1 4.6 5  7.65 11.21 11.35 9.86 11.27  j  j  9 9 9 9 12 12  4.8 5.6 6.6 5.9 4.5 8.6  12 12  9.4 7.6  0.54 1.93 2 . 23 2.22 1.73  5 5  c 3  3 3  4.4 4.0 • 5. 9 4.6 4. 1 4.0  4.40 4.59  3. 39 4.64 1.03 8.03 7.20 6.15  -  27.1 3 2 . 3 44.4 5.3 5.3 6.2  1. 1 0.4  D.  | j |  M  s  10 10 10 10 10  GRAZING PEEIOD (t=11,34)  2.36 1.95 5.66 3 . 17 2 . 12 2.65  9  I I  9 9 11 11  [  11  5.8 6. 1 €.3 5.5 5.7 6.8  2.11  3.f  2 4.37 2.29 3.32 •1.51  113 Table  4.  R e s u l t s o f t h e o y s t e r growth d u r i n g E x p e r i m e n t 2 1 . GRWM and GR1S, r e f e r t o t h e growth r a t e o f t h e meat and s h e l l w e i g h t s per o y s t e r p e r week f o r a c o m p a r i s o n w i t h EXP 3 A and EXP .5. MSB ATI C i s t h e r a t i o between NETWM and NETWS.  SURF ACE  DEPTH  SUESTN  NETWT  NETWM  PERWK GRHM  1 2 3 4  81.0 88.5 55.5 65. 8  11.0 19.5 14.0 14.0  17.3 26.0 22.4 25.9  0. 16 0.24 0.23 0.28  NETWS  GBWS  MSEATIO  70.0 65.0 41. 0 51.5  1.00 G.€6 0.68 1.03  . 162 .283 .311 .272  NETWS  GRWS  MSEATIO  5 7.0 64,5 55.5 65.5  0.81 0.76 0.74 0,80  . 202 .364 .405 .244  NETWS  GliWS  MSRATIG  4 0.0 8 8,0 74.0 29.5  0.73 1.17 0.87 0.54  .313 .267 .324 .356  .MID DEPTH SUESTN 1 2 3 4  NET1T  NETWM  PIEWM GRWM  68.5 88.0 78.0 81.5  11.5 23.5 22.5 16,0  11.6 36.4 36.6 15. 2  EOTTCP SUESTN 1 2 3 4  NETWT 52.5 111.5 98.0 40. 0  0.16 0. 28 0.30 0.25  DEPTH  NETWM  PEEWM GRWM  12.5 23, 5 24.0 10.5  14, 1 25,1 23.3 10.9  0.23 0.31 0.28 0. 19  114  Table  5. Growth o f t h e s c a l l o p s d u r i n g E x p e r i m e n t 2B, See A p p e n d i x 2 f o r an e x p l a n a t i o n o f t h e variables.  MID STATION SOESTN 1 •2  3 4  NSUBV 4/8 4/8 3/8 3/8  I 5. 0 5. 6 6. 1 6. 8  SD  PEEL  .58 0.5 . 19 - 1.0 .35 -0.1 .38 -0.2  W 4.4 5.1 5.4 6.2  .PEHW  1GTT  SD  2, 0 .58 0. 1 .20 1.7 .32 .44 -0.5  16. 1 23,5 29.8 43.8  4.36 0.51 . 3.33 6.62  -0.6 -1,7  SD  PER ET  SD  PEBRT -0.2 1.1  BOTTOM STATION SOBSUN NSURV 5 8 9 12  8/8 8/8 8/8 8/8  L  SD  FEE I  4.7 .59 0.3 5. 6 .27 0.1 6. 1 . 22 0.0 6.7 .35 -0.3  4.2 4.9 5.6 6.1  SD  PEEW  SGTT  .62 .25 .21 .43  0.7 0.3 0.3 0.0  13.8 22.0 32.5 44. 3  -9.6 4.11 3,10 -6.S 3.77 -5.0 6.76 -10 . 5  115  T a b l e 6.  D e s c r i p t i v e s t a t i s t i c s summary o f s e l e c t e d v a r i a b l e s f o r EXP3Aj i n c l u d i n g a breakdown i n t o t h e p r e - g r a z i n g and g r a z i n g p e r i o d . See a p p e n d i x 1 f c r a d e f i n i t i o n o f t h e v a r i a b l e names.  TOTAL EXPERIMENT (t=1,21) VAR  | STN I  SR TEMP  | l J I I I i i i f t t i I I I I I I  I I S M B o I I s M E o i s M  !  B  NO 3 NB3  OXY  CHLA  PROD ASS  B  o s M  I "0 I s I M I B i s | M i E  N  21 I 21 I 21 |21 | 21 I 21 I 21 I 21 I 6 I 6 I 6 I 21 I 21 I 21 I 21 I 21 I 21 I 13 I 13 1 13 | 13 i 9 I 9 i 9 \ 9 I 9 | 9  MEAN 330. 11.6 16.3 16.2 16.2 16.2 21.0 0.50 0.40 0.34 0.27 0.50 7. 04 9.91 9. 88 9.97 9. 9 2 10.4 11. 1 11.5 1 1. 2 25.0 29.3 27.2 3.4 3.8 3. 9  SD  PRE-GRAZING (t=1,6) |  N  MEAN  104. | 1.36 I G.S6 | G.92 | 0.89 | C.98 | 2.05 | 0.28 J 0.29 | 0.31 J 0.17 J 0.50 | 0.72 J 1.89 \ 1.9 1 | 2.51 | 1.69 | 9.7 4 J 11.20 j 11.42 | 11.50 | 19.09 \ 2 1.94 | 16.83 I 1.98 | 2.33 | 2.71 |  6 6 6 6 6 6 6  400. 11.1 15.9 15.8 15.8 15.8  6 6 6 6 6 6 6 6 6  6.73 10.50. 10.56 10.54 10.36 13.9 15.8 16.2 16.0  3 3 3  2.3 2.8 2.2  20.4  SB  GRAZING PEEIOD Jt=7,21) J  55. .j 0. 17 | 0.49 | 0.54 | 0*49 | 0.46 | 1.35' |  0.13 2. 84 2.£€ 2.84 2.62 12.7G 14.80 14.99 15.10  N . Ml AN 15 15 15 15 15 15 * .  | j |  15 15 15 15 15 7 7 7 7  C.55 ; 1.80 1.23 |  6 6 6  J 1 |  300. 11.8 16.4 16.4 16.3  16.4  SD 108. 1.58 1.08 1.00 0.9 8 1.09  N/ft  7.16 9.68 9.60 9.74 9.74 7.3 7.2 7.5 7.1  €.78 1.41 1.40 1.75 1.23 5.69 5.34 5.78 5.64  4.0 2 .18 4.3 2.55 4.7 2.S4  116 Table  7.  J e s u i t s o f t h e o y s t e r growth d u r i n g E x p e r i m e n t 3 f l . GBWK and G.BHS r e f e r t o t h e growth r a t e c f t h e meat and s h e l l w e i g h t s p e r o y s t e r p e r seek f o r a c o m p a r i s o n w i t h EXP2A and EXP 5. MSB il TIC i s t h e r a t i o between KET«H and NETWS.  SUBFACE SUESTN 1 2 3 4 5 6 7 8  DEPTH  NET AT  NETWM PIBWM  GBSM  18.5 9.5 8.0 16.5 16.5 6.5 24.5 28.0  4,8 2.0 2,0 2.5 0.5 1.0 11.0 5.0  0. 12 0.06 0.08 0.13 0.02 0.03 0.61 0.21  4.4 1.4 1.9 3.8 C.8 2.6 11.2 «f.3  MSBftT I G  NETHS  GBHS  14.5 7.5 6.0 14.0 16.0 5.5 13.5 23.0  0.45 0.25 0.25 C.7C 0.53 G.20 0.15 0.S6  .276 .267 .333 .179 .031 .182 .815 .217  NET1S  GESS  MSBSTIC  4.0 17.0 10.0 1.0 7.0 7.5 7.5 6.5  0.20 0.50 0.46 0.05 0.27 0.47 0.31 0.27  .500 . 147 .200 1.000 .643 .067 .267 .154  NETWS  GBSS  MSEATIG  C.92 0.17 0.69 0.77 1. 19 0.06 0.50 0,47  .202 .727 .12 1 .304 .256 1.000 .44 1 .667  MID DEPTH SUESTN  NET ST  1 2 3 4 5 6 7 8  6.5 19.5 12.0 2.C 11.5 8.0 9.5 7.5  NETS M IE EH B GBSM 2.0 2.5 2.0 1.0 4.5 0.5 2.0 1.0  3.5 1.8 3.3 2.4 7.2 0.8 2.9 1.5  0. 10 0,08 0.0 9 0.05 0. 18 0.03 0.08 0.04  BOTTOM STATION SUESTN 1 2  •3  4 • 5 6 7 8  NET8T  NETWM PEBH M GB8M  31.0 9.5 18.5 30.0 27.0 4.0 24,5 25.0  5.2 4.0 2.0 7.0 5.5 2.0 7.5 10.0  4.4 6.5 3.5 10.9 7.1 2. 2 4.8 10.6  0. 18 0. 12 0.08 0.23 0.30 0.06 0.22 0.6 2  25.8 5.5 16.5 23.0 21.5 2.0 17.0 15.0  117 facie  8.  D e s c r i p t i v e s t a t i s t i c s summary o f s e l e c t e d v a r i a b l e s . f o r EXP3B, i n c l u d i n g a breakdown i n t o t h e p i e - g r a z i n g and g r a z i n g p e r i o d . See Appendix 1 f o r a d e f i n i t i o n o f t h e v a r i a b l e names. .  TOTAL EXPERIMENT (t=1,34) V8R  I STN ,1  SB TEMP  I J 1 ] J 1 1 I'  I I s M B 0 I S  1  E o I s  NG3  i  NH3  1  1 i 1 1 i  OXY  CHLA  1 1 1 1 1 1 ] I i  EIOD  1  ASS  1 1 I 1 j  N  MEAN  SD  J  N  MEAN  SD  1 34 1 34 34 1 34 | 34 1 34 1 34  380. 11.4 14.9 14.8 14.8 14.8 19.3  105. G.52 1.05 1.04 1.04 1. 10 2.02  \ | | J | | | J  11 1 1 11 1 1 11 11 11  410. 11.5 15.5 15.5 15.5 15.5 18.9  102. 0.32 0.78 0.7 6 0.74 0.63 1.26  B  11 11 11 11 1 1 9 . 9 9 9  6.93 10.39 10.44 10.42 10.41 14. 9 15.8 15.2 16.2  4.4 4.3 4.4  GRAZING EEE.IOD (t=12,34) N  MEAN  SD  23 23 23 23 23 23 23 23 23 23 23  366. 11.4 14,5 14.5 14.5 14.5 19.6 0.3 0.3 0.3 0.2  106. 0.59 1.02 1.02 1.01 • 1.06 2.29 C.37 0.34 0.38 0.3 2  0.2 1 2. 14 2. 18 2. 16 2.05 j 13.70 | 14. 48 14.14 j 15.06  23 23 23 23 23 9 9 9 9  7.01 12.01 12.03 11. S6 11.7.4 29.2 29.7 2S.9 29.7  2.51 2,5 9 j 2.33  9 9 9  | „i  | | | |  I  | |  H'  E o I s M E o s M E o s M E s H B  PBE-GBAZING <t=1,11)  I 1 I | | | | I | | 1 | | ] I | I | I 1  34 11 11 11 34 34 34 34 34 34 18 18 18 18 14 14 14 14 14 14  0.42 0.59 0.61 0.60 0.48 6. 98 11.49 11.52 11. 46 11.31 22. 1 22.8 22.6 23.0 38.3 36.9 39.8 2.4 2.2 2.3  0.29 J 0.47 J G. 35 | 0.31 | 0. 18 I 0.54 J 1.59 | 1.58 | 1.54 J 1.40 | 12.37 | 12.54 | 12.55 | 12.78 | 13,86 | 14.98 | 19.79 J 2.11 | 2.13 | 2.13 |  5 5 5  C.65  o.so  0.€4 0.79 0.66 4.67 4.08 . 3.56 4.20  1.2 0.28 1.1 0.10 1.0 0.26  T a b l e 9. Growth c f t h e s c a l l o p s d u r i n g E x p e r i m e n t See Appendix 2 f o r an e x p l a n a t i o n c f t h e variables.  SURFACE STATION SOBSTN NSUBV 6 7 10 11  0/8 0/8 0/8 0/8  L  SD  3.6 4, 3 5.4 5.8  .23 .39 . 19 .44  PES I N/a N/A N/A N/A  M  SD  3.1 3.8 4.8 5.2  PEBW  .19 N/fl .41 . N/A .17 N/A . D2 N/A  KGTT 6.0 10.6 19.4 26.3  SB  PEE ST  1.08 . N/A 2.89 N/A 1.27 N/A 4.83 N/A  MID STATION SUESTN  NSUBV  1  1 2  3/8 4/8 6/8 5/8  3.6 4„2 5.2 5.9  3 a  SD  PEEL  0.3 .22 .29 0.0 .31 0.3 .53 '• G.2  BOTTOM SUESTN 8 5 9 12  NSUBV  L  7/8 8/8 8/8 8/8  3„8 4.5 5. 1 5.7  SD .45 .27 .15 .31  SD  PEEW  KGTT  .19 .19 .32 .51  -0. 1 0.1 1.6 0.2  6. 2 S.8 18.3 26.6  « 3.1 3.7 4.7 5.3  SD 1. 12 1.35 2.48 6.92  PEEST -0.4 -1.9 1.6 0.8  STATION  PEEL  H  1.8 1.0 0.4 0.4  3.3 t).0 4.6 5. 1  SD  PERW  .42 .26 .20 .34  0.5 1.0 0.3 0.5  KGTT 6.3 11.4 16.9 2 5.4  SD 1.82 1.11 2.87 4.23  PES«T 11.7 14.4 16.8 1.2  119  • fable  EXP 4 A  10.  P e a r s o n c o r r e l a t i o n c o e f f i c i e n t s between t h e f o r c i n g v a r i a b l e s during Experiment 4 f o r bcth Tank a (upper r i g h t ) and Tank E (lower l e f t ) . The v a r i a b l e s i n c l u d e : i n c i d e n t s o l a r r a d i a t i o n , SB ( l y / d a y ) ; i n f l o w t e m p e r a t u r e , TEMP ( d e g - C . ) ; , i n f l o w n i t r a t e c o n c e n t r a t i o n , K03 ( u E / l i t r e ) ; and i n f l o w oxygen c o n c e n t r a t i c n , OXY ( n g / 1 i t r e ) . As t h e p r e f i x A i n t h e v a r i a b l e name i n d i c a t e s , t h e v a r i a b l e s were ' f i r s t a v e r a g e d ' , s c t h a t AVAE = 0.5*(VABt + V'fiBt+1), f o r a c o m p a r i s o n with the d a i l y i n t e g r a t e d s c l a r r a d i a t i o n , S B . SB h a s been l a g g e d one t o t h r e e days (SB1 t o SB3) t o examine t h e s e r i a l c o r r e l a t i o n f o r solar radiation.  SB  ATEMP  A NO 3  AOXY  .065 t=33 S=,.6 40  .345 t=33 S=.050  -.769 t=33 S=.0O2  .678 t=33 £=.002  SB1  SB 2  SB3  v A  E 4 E  . 118 t=3 3 S=.512  SE  A TEMP  .080 t = 33 S=.660  AN03  .008 t=33 S=.964  -.844 t=3 3 S=.002  AOXY  .356 t=33 S=.042  .662 t=33 S=.002  SE 1  .422 t=32 £-.016  -. 13 7 t=3 2 S=.254  SB 2  -.118 t = 31 S=.528  -.33 2 t=31 S=.068  SB3  -.433 t=30 S=.016  -.307 . t =30 S=.098  -.598 t=33 £=.002 -.657 t=33 S=.C02  .422 f=32 • S=.C16  -.118 t=31 £=.528  -.433 t=3C £=.0 16  -. 137 t=32 S=.456  -.364 t = 31 S=,044  -.365 t=3 0 £=.0 48  120 Table 11,  D e s c r i p t i v e s t a t i s t i c s of .'selected,, variables' f o r 1XP4A, i n c l u d i n g s t a t i s t i c s f o r the n i t r a t e depleted p e r i o d ( t = 2 9 ) . See appendix 1 f o r the d e f i n i t i o n of the v a r i a b l e -names,,. T  Ml. AN  Sa D.  34 34 34 34 34 34 B 0 34 TEMPTS S 34 M 34 B 34 34 0 34 I WO 3 . OXY I 34 : s . 34 M 34 B 34 34 0 OXYN S 34 M . 34 B 34 34 0 saT 34 I S 34 E 34 B 34 0 34 CHLfl S 34 a 34 34 .B 0 34 29 SB i TEMP 29 i S 29 M 29 B 29 0 29 s 29 TEMP N M 29 B 29 0 29 N03 .1 29 N03N s 29 M 29 B 29 1 o 29 29 OXY I s 29 29 H  378, 27.3 10.4 14.7 14. 6 13. 1 14.6 4.3 4.2 2. 7 4.2 18.8 7.58 10.85 10.90 8.98 10,69 3.26 3.32 1.39 3.11 81. 126. 127. 102. 125. 15.4 15.4 18. 2 16.3 358, 10.4 14.7 14.6 12.8 14.6 4.3 4.2 2.4 4.3 19.0 18.7 18.S 15.3 18. 1 7.47 11,20 11.30  149.7 0.79 0.41 1.29 1.2 6 1.54 1.30 1.32 1.28 1.41 1.3 2 2.56 0.682 1.364 1.406 1. 163 1. 179 1. 824 1. 822 1, 06 2 1.691 7.6 17.7 18.1 15,2 18.8 7.20 7.09 8.82 7.82 149,0 0.39 1.27 1.23 1.39 1.27 1.28 1.24 1.30 1.28 2.6 5 3.06 2.81 5.56 2.98 0.669 1. 14 2 1. 112  van SH SAL TEMP  STN I I I S M  S . E . ., C.v. E A NGE  Max  25.7 3S.6 453. . 564, 3.0 26.5 0. 14 2.9 0,07 3,9 1.3 11.1 0.22 4.1 1.6.6 8.8 0.22 €.6 4.1 16.3 0.26 11.8 5.1 16. 1 0.22 4.4 €.9 16.5 0. 23 30.7 '.. 4.9 •6. 1 0.22 30.5 4.9 6.1 0.24 52.2 4.9 5.3 .0, 23 31.4 .' ,5.0 6.1 0.44 13.6 9.5 22.2 0.117 S.C 3.23 9.S7 ,0.234 12.5 5.04 13.26 0.241 12. $ 5.20 13.69 0. 199 12.9 ' 5.03 11.01 0.25 4 13.8 5.S5- 13.28 56.0 6.40 6.21 0.313 0.312 5 1 . S 6.73 6.81 0. 182 76.4 4.34 2.S 1 0,324 6 0 . 8 6.53 6.10 107. 1.3 S.4 35. 3.0 14.0 65. 160. 3.1 14.3 66. 163. 2.6 14.S 6 8. 1 3 1 . 3.2 15.0 74. 160. 1.23 46.6 28.6 28.9 1.22 46.0 25.3' 2 5.4 1.51 48.5 33.1 33.2 30.7 1.34 48.0 3 0.8 27.7 11.6 445. . 55'6. 0.07 3.8 1.3 1-1.1 0.24 8.6 4.0 16.6 0.23 8.4 1& 3.7 0.26 16.1 10.8 5.1 8.7 0.24 3-9 16.5 0,24 2S.8 4.1 €.1 0. 23 29.5 4. 1 €. 1 0.24 54.2 4.9 5.3 0.24 29.8 4. 1 t. 1 0.49 13. 9 9.5 22.2 0.57 16.4 10.5 22.2 0.52 14, S 10.6 2 2.2 1.03 36. 3 24.5 22. 1 0.55 16.5 13.1 22.2 0. 124 • S.O 3.23 . 9 . 97 0.212 10.2 3.76 13 .26 0.207 S.8 4. 08 13.69  ;12'1 29 29 29 29 !' B • 29 ! o 29 I I 29 s 29 I H 29 ; B 29 0 29 l " s 29 r M 29 29 B I o 29 j s 29 I • M 29 B 29 I o 29 29 I 's I .fit 29 29 B I . .0 29 29 I. s | H 29 B 29 I , 0 29 1 I 29 |. S ! H 29 ! B 29 I ' o. 29 1 I.I 29 I s I n 29 29 B I o 29 1 I t • s 29 I. , M 29 I E 29 I o 29 1 I I j ' s 29 I H 29 i B • 29 I o 29 Ii : 1 l s 29 I H 29 29 i B I o 29 1 I I i s 29 | B 29 I • B 29 I o 29 I  OXYN  SAT  CHLA  CHLE  CHLC  Cl  EA  CA  CTA  EC  ECT  CCT  B.  0 • s  8,93  11,04 3,73 3.83 1.53 3.5 8 79. 131. 132. 101. 129, 17.2 17. 2 20.5 18. 3 .1.4 1.4 2.3 1. 4 11.0 12.2 12.1 11. 1 21.8 2 1.6 2 3.2 23.2 . 105 .092 .086 .119 .081 1. 153 .660 .634 .598 .624 1.123 1. 283 1.277 1. 153 1.274 . OS 1 .136 .13 2 .200 . 127 .093 .073 , . 06 8 . 102 . 065 1.027 .526 .510 .521 .486  1. 251 1.306 1.5.33 1.439 1.080 1.632 7.4 15.6 1-5.3 16.2 17.2 5.37 5. 16 6.74 5.90 0.70 0.80 1.18 0.80 2.93 3.4 9 4.01 2. 97 6.76 6.00 7,08 7.26  0.232 0.243 0.285 0.267 0. 201 0.303 1.4 2.9 2.8 3.0 3.2 1.00 0.S6 1.25 1.10 0. 13 0, 15 0.22 0.15 0.54 0.65 0.74 0.55 1.26 1. 11 1.32 1.35  .049 .051 .014 .046  .009 .010 .014 .009  53.3 .188 5 9 . 3 .202 62.2 : .395 56.8 .201  .105 .115 .103 .057  .020 .021 .019 .018  15,9 18.1 17.2 15.5  .35 8 .405 .402 .481  .652 . ES3 .168 .943  .209 .196 .15 2 -159  .039 .036 .028 .030  16.3 15.3 13,2 12.5  .768 .677 .599 .573  1.127 1. €09 1.5(17 1. 590  .065 .013 .112 .065  .0.12 .014 .021 . 012  47.8 .257 .255 55.3 56.0 -•..553 .207 51.2  . 267 .259 . 561 .223  ,038 .038 .055 .037  .007 .007 .010 .007  52.1 55.9 53. $ 56. 9  .140 . 135 . 293 .16 8  .110 .128 .072 .057  .021 .024 .013 ,018  20.9 .45 4 2 5 . 1 • .559 13.8 .301 19.6 .459  13..S 11.8 41.1 37.6 10.6 45.6 9.3 11.9 11,6 16.0 13.3 31.2 3 0,0 3 2-9 32.2 50.0 51.1 51. 3 51. 1 26.6 28.6 3 3. 1 26.8 31.0 27.8 30.5 31.3  5.03 11.01 5.95 13.28 6.40 6.21 5.69 6.8 1 4.34 2.91 6.53 6.40 35. 101. 160. 52. • 55. 163. 68. 131. 74. 160. 2 2 . 1 2 8.9 1 6 . S 2 5.4 23.6 3 3 . 2 22.6 30.8 2.6 5.7 3.2 3.2 4.6 5.2 3.2 3.4 12.4 18.2 14.4 20.7 13.6 18.0 12.1 11.2 24.8 35.8 19.9 3 1.6 29.8 4 2.5 27.2 31,0  .135 . 135 .288 . 161  .154 a 20 2 .400 .210  .119 .£62 .646 .154  122 Table  12,  Descriptive s t a t i s t i c s of selected variables f o r FXP4E, i n c l u d i n g s t a t i s t i c s f o r t h e n i t r a t e d e p l e t e d p e r i o d ( t = 2 7 ) . See Appendix 1 f o r t h e d e f i n i t i o n of t h e v a r i a b l e canes.  VfiB  STN  SB SAL TEMP  34 34 34 34 34 34 34 34 34 34 34 34 34 34 34 34 B 34 0 34 S 34 H 34 B 34 0 34 I s 34 34 M 34 B 0 34 . s 34 34 M 34 B o 34 I 27 I 27 ' s 27 M 27 B 27 ' '0 27 s 27 27 M 27 B o 27 27 i s 27 27 M 27 B o 27 I 27 s 27 27 M  TEH FN I N03 OXY  OXYN  SAT  I I  CHLA  |I  I I SB TEMP  I  I  I |  I TEMFN  I I  I I  NQ3 N03N  OXY  I  i  I I I  1 | 1 1  I I I S M B • 0 S M B 0 I I S M  T  MI AN  S.D.  378. 149.7 0.79 27.3 10.4 0.39 13.0 0.78 13.0 0.76 0.76 13.0 13.0 0.78 0.76 2.7 2.6 0.14 2.6 0.76 2.6 0.76 18.7 2.87 7. 55 0.670 11.58 1.645 11.48 1.653 1 1.28 1.679 11.27 1. 487 4.03 1.904 3. S3 1.903 3.70 1.835 3.71 1.711 80. 7.4 . 131. 18.9 130. 19.0 127. 19.0 127. 17.2 31.5 15.93 33. 1 16.84 34.0 17.09 33.1 16.97 3 45. 146.4 0.40 10.3 12. 9 0.76 12.9 0.74 12.9 0,76 12.8 0.75 2.6 0.76 2.5 0.75 2.5 0.78 0.75 2.5 19. 1 2.82 19. 1 2.81 19. 1 2,82 19. 1 2.81 19. 1 2.79 7.39 0.644 12.26 1.022 12.13 1. 121  S. E. , C.V. BANG! 25.7 0.14 0.07 0. 13 0.13 0.13 0.13 0. 13 0.13 0. 13 0.13 0.46 0.115 0. 282 0.284 0.288 0.255 0.327 0. 326 0.315 0.293 1.3 3.2 3.3 3.2 2.9 2.73 2.89 2.93 2.91 28.2 0.08 0.15 0.14 0. 14 0. 14 0. 15 0.14 0. 15 0. 14 0.54 0.54 0.54 0.54 0.54 0. 124 0. 187 0.216  39.6 2.9  3.8  6.0 5.8 5.S 6.0 28. 1 28.5 29.2 29.2 15. 3  e.s  14.2 14.4 14. 9 13.2 47.2 48.4 49.6 46.1 9.2 14.4 14.6 15.0 13.5 50.6 50.9 50.3 51.3 12.1 3.9 5.9 5.7 c c —• • > 5.9 28.5 30.0 31.2 30.0 14.8 14.7 14.6 14.7 14.6 8.7 8.3 9.2  453. 3.0 1.3 2.4 2.4 2.5 2.5 2.7 2.6 2.6 2.6 11.3 3.23 5.61 5.41 5.72 5.2 0 6.7S 6.60 5.92 5.66 36. 67. 70. 61. 67. 51.6 54. 3 54.1 52.4 445.  1.3  2.4 2.4 2.5 2.4 2.6 2.6 2.6 2.5 11.3  fi 2 X 561. 2 6.5 11.1 14.2 14.2 14.2 14.3  3.7 3.6 3.6 3.8  2 2.7 9.S7 14. 13 13.S6 14.30 13.75 6.67 6.74 6.02 6.00 107. 164. 1€5. 158. 165. 51.7 54.4 51.2 5 2.6 5 56. 11. 1 14.2 14.2 11.4 11.2  3.6 3.6 3.6 3.5  2 2.7 22.7 2 2.7 11.3 2 2.7 11.3 22.7 11.3 9.S7 3.23 3,87 14. 13 4. 21 13. 56  11.3  12 3  I OXYN  SflT  CHLA  CHLE  CHIC  CT  EA  Cfl  CTA  EC  ECT  CCT  B 0 S  27 27 27 H 27 |.: E 27 I o 27 i I 27 s 27 I M 27 I 3 27 I o 27 i s 27 I M 27 | B 27 | 0 27 I s 27 | CS 27 I B 27 I- o 27 I s 27 I H 27 I B 27 I o 27 I • s-27 I a 27 ! B 27 I o 27 | I 1 |- s 27 I M 27 I B 27 I o 27 I I 1 I s 27 I M 27 I B 27 I o 27 I I 1 i s 27 I H 27 i E 27 1 o 27 1 I 1 I s 27 I fi 27 1 B 27 1 o 27 1 I 1 1 s 27 | M 27 I B 27 i o 27 | I 1 1 s 27 1 M 27 1 B 27 1 o 27  11.81 11.86 4. 87 a. 74 4.45 4.47 79. 138. 137. 133. 134. 38.8 40. 8 41.8 40.6 1.7 1.5 1.6 1.6 20.2 20. 0 20.8 20. 6 43.0 45.2 45.7 46.0 . 105 .047 .039 .040 .041 1, 153 .528 .497 ,503 .519 1.123 1 , 120 1.118 1. 104 1.138 .091 .086 .075 .078 .07 4 .093 .045 .037 .037 .036 1.027 .481 . 453 .459 ,461  1.336 0.S70 0. 935 1.045 1. 111 0.806 7. 1 13.0 13.9 15.8 12.4 7.08 7.57 7.46 7,73 1,50 1. 11 1.28 1. 46 3.19 3.02 3.49 3 . 15 7.70 7.84 6.94 7-78  0 . 257 0 . 187 0. 180 0.201 0.22G 0. 155 1.4 2.5 2.7 3.0 2.4 1.36 1.46 1.44 1.49 0.29 0.21 0.25 0.28 0.62 0.58 0.67 0.61 1.48 1.51 1.34 1.50  11.3 8. 2 19.2 22.0 25.6 18.0 9.1 9.4 10.1 11.9 S,3 18.2 18.6 17.8 18.S 88.2 74.0 80.0 91.3 15.8 15. 1 16.8 15.1 17.9 17.3 15.2 16.9  4 . 9 S 14.30 3.93 13.15 3.43 6.67 4 . 2 2 6 .7 4 3.7S 6.02 3.0 0 6.00 36. 107. 50. 164. 57. 16 5 . 5 5 . . 158. 56. 165. 27.2 5 1 . 7 25.0 54.4 26.6 51.2 30.4 52.6 5.9 5. 9 4.4 4.4 5.4 5.4 4.S 1.9 13.4 2 6 . 5 11.3 24.6 13.8 2 5 . 9 11.2 25.6 34.4 5 6 . 1 31.3 5 6 . 9 27.6 59.5 35.6 5 5 . 8  .044 .03 5 .033 .04 3  ,008 .007 .006 .008  93.6 89.7 82.5 104. ,  .179 .118 .148 .136  .066 .065 .066 .073  .013 . 012 .013 .014  12. 5 13. 1 13. 1 14.1  . 2 7 5 . 681 .284 . 68 9 . 2 9 0 . 669 m 28S .707  . 168 .161 . 133 .145  .032 .030 ,026 .028  15.0 14.4 12.0 12,7  .706 .659 .56 9 .651  .078 .063 .065 .070  .015 .012 .012 .014  90.7 64.0 83.3 S4.6  * 3 3 3 ..333  .291 .285 .255  .291 .28 5 .255  .046 . 036 . 03 2 .038  .008 .007 .006 .007  102. 97. 3 100. 106.  .186 .147 .140 .131  .166 . 147 .140 . 13 1  .087 .093 .064 .076  .017 .018 .012 .015  18.1 • . 3 9 9 2 0 . 5 .437 13.9 . 2 7 3 ie.5 .377  . . . .  179 148 148 136  1,5 44 1.5 28 1.509 1. 562  .708 . 7 39 . 569 .719  Table  13.  R e s u l t s o f t h e a n a l y s i s o f v a r i a n c e and m u l t i p l e c l a s s i f i c a t i o n a n a l y s i s f o r s e l e c t e d v a r i a b l e s f o r EXP4A a s a f u n c t i o n o f t h e i n d e p e n d e n t f a c t o r s TIME and STATION. F v a l u e s and s i g n i f i c a n c e l e v e l s i n t h e ANOVA a r e p r e s e n t e d f o r t h e n i t r a t e - d e p l e t e d period only, s i n c e the s i g n i f i c a n c e values f o r the t o t a l e x p e r i ment (t=34) a r e s i m i l a r . The t e n s a m p l i n g t i m e s f o r t h e p r o d u c t i v i t y v a r i a b l e s i n c l u d e e v e r y t h i r d day f r o m Day 6. D a t a from t h e b o t t o m s t a t i o n h a s been e x c l u d e d (N/I) i n t h e a n a l y s e s o f t h e o t h e r v a r i a b l e s (TEMP t o CCT)..MULT R i s t h e m u l t i p l e c o r r e l a t i o n between t h e d e p e n d e n t v a r i a b l e and b o t h i n d e p e n d e n t v a r i a b l e s TIME and STATION. The MCA t a b l e i n d i c a t e s t h e e f f e c t o f each c a t e g o r y o f STATION, e x p r e s s e d a s a d e v i a t i o n f r o m t h e g r a n d mean, and shows t h e t i m e o f t h e minimum and maximum d e v i a t i o n s d u r i n g t h e n i t r a t e - d e p l e t e d period.  V AR NAME  1 ]  TEMP J TEMPN J OXY J OXYN i SAT | CHLA | CHLB | CHLC | CT | BA | CA J CTA J BC J BCT \ CCT | PGO | PNO | RES J PBOD J PGODY \ PCDY J PGOST J PNOST | RESST I ASS J EXCST ] ALPHAG] ALPHAC]  ANALYSIS BY T I M E F T SIG. 29 29 29 29 29 29 29 29 29 29 29 29 29 29 29 10 10 10 10 10 10 10 10 10 10 5 10 10  *** ***  16. 27. 2216. 14. 18. 8. 1 16. 3-3 12. 1617. 3.6 2.8 2-7 5.6 2. 1 3.6 2.2 5.0 2. 7 5-0 4.0 6-8 13. 24.  .000 .00 0 .000 .000 .000 .000 -000 -000 .000 -000 .000 -000 .000 -000 .000 -030 -036 .00 1 -092 .011 -079 .002 .034 .002 .006 .011 .000 .000  OP V A R I A N C E | BY STATION ] A F SIG. | 3 J 3 | 3 J 3 | 3 j 3 ] 3 | 3 ] 3 ] 3 I 3 J 3 i 3 J 3 ] 3 ] 3 | 3 ] 3 J 3 | 3 ] 3 j 3 ] 3 | 3 ] 3 ] 3 3 3 . 3  8. 5 8.5 1.9 1. 9 1.7 2.4 0. 6 0. 8 1. 8 2. 2 1.5 0, 1 0. 8 2-2 0.9 1. 4 1.3 0. 0 5. 4 1- 2 5- 1 2. 1 0.5 0. 9 0. 6 7.3 5.6 16.  | MULT J GRAND | ] MEAN ] I . S  -00 1 .001 | . 16 % \ .161 I .185 j .096 j -56 7 | -447 j .178 | . 119 -224 j -929 j .479 j -115 | .409 | .274 \ -287 | .984 j .014 j .338 .018 i . 153 I .600 j .435 | .588 | .015 ] .013 J .000 |  .998 .998 .942 -965 .957 . 942 -937 .949 . 897 .942 -793 -928 . 944 .947 .803 -780 -773 .859 - 787 .810 . 788 - 856 -766 - 850 -821 -916 -935 .966  ] |  MULTIPLE CLASSIFICATION ANALYSIS DEV'N BY S T A T I O N | D E V N BY T I M E SUR MID OUT | BOT M I N . DAY ] MAX.  14.6 J .05 4-3 j . 05 I 11.18 ] -02 I 3.71 | - 02 I 130. 1 -3 117- 6 ] -.38 | 1.4 | .05 ] 11.0 ] . 06 | 22.2 1 -0.4 ] .09 1 .01 1 .64 ] .02 ] 1.28 ] -00 | 0. 1 3 ] .00 ] 0.07 ] .00 | 0.51 ] .02 | 220. I " 1 1 . | 181. | -12. I 40- I 1] 38. 1 -63 2. 11 J . - - 0 9 \ 0.35 J - . 0 5 I 11-6 ] 0.9 ] 9.2 ] 0.6 ] 2.4 | 0-4 ] 2.0 J - 0 . 1 1 6-5 j 0.4 I 6 3 - 2 1-10.0 \ 11.6 ] - 2.8  -.06 -.06 . 12 -12 1.2 -.38 -.04 -.23 -0-6 .00 .00 .00 .00 .00 .00 -18. -17. 0-5. -- 1 6 --04 0- 1 0-0 0. 1 -0. 1 1.5 -1.6 -0.7  N/I N/I N/I N/I N/I N/I N/I N/I N/I N/I N/I N/I N/I N/I N/I 29. 29 -11 1.25 1.00 -1- 1 -0.6 -0.5 0. 1 -2.0 11.6 3.5  -0 1 -01 13 --13 -1-5 .76 .00 - 16 1.0 -.01 -.0 2 -00 .00 .00 -.01 N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A  | -2.03 ;| - 2 . 2 7 j| - 1 . 4 9 - 3 . 45 \ -22. 6 -8. 87 -1.23 \ -4.39 j -10.9 -0. 08 | -0. 13 | -0- 25 - 0 . 11 -0. 06 | - 0 . 16 -115. j - 9 3! -22. -14j -1-0 1 ] - 0 . 15 | - 4.3 i - 3. 3 ; - 1 . 3 | - 0.6 | - 3. 6 • -23. 6 | - 7.0  24 31 30,31 13 30 13 20 14 28 20,25 25 29 20,25 20,25 20 30 30 30 9 30 9 30 30 24 24 30 15, 18 12  I | J 1 ] ] ] 1 J ] I | ] | i ] ] | I ] ] | I | | 1 ] ]  DAY  1-84 34 1-83 19,20 19 2- 1 7 2.76 20 29. 6 19 9.70 24 1.41 6 6.87 6 9-1 7 34 .10 28 -19 19 .31 .12 34 .06 28,34 9 - 15 86. 6,21 102. 6 65. 12 20. 27 6 1.11 0.16 6 12 4.3 4. 1 21 5- 4 12 34 1. 0 5 1.7 48. 1 21 30 17.5  Table  14.  R e s u l t s o f t h e a n a l y s i s o f v a r i a n c e and m u l t i p l e c l a s s i f i c a t i o n a n a l y s i s f o r s e l e c t e d v a r i a b l e s f o r EXP4B a s a f u n c t i o n o f t h e i n d e p e n d e n t f a c t o r s TIME and STATION. F v a l u e s and s i g n i f i c a n c e l e v e l s i n t h e AN0VA a r e p r e s e n t e d f o r t h e n i t r a t e - d e p l e t e d period only, since the s i g n i f i c a n c e values f o r the t o t a l e x p e r i ment (t=34) a r e s i m i l a r . The t e n s a m p l i n g t i m e s f o r t h e p r o d u c t i v i t y v a r i a b l e s i n c l u d e e v e r y t h i r d day f r o m Day 6. Data from t h e bottom s t a t i o n has been e x c l u d e d (N/I) i n t h e a n a l y s e s o f t h e o t h e r v a r i a b l e s (TEMP t o CCT) . MULT R i s t h e m u l t i p l e c o r r e l a t i o a between t h e d e p e n d e n t v a r i a b l e and b o t h i a d e p e n d e n t v a r i a b l e s TIME and STATION. The MCA t a b l e i n d i c a t e s t h e e f f e c t o f e a c h c a t e g o r y o f STATION e x p r e s s e d a s a d e v i a t i o n f r o m t h e g r a n d mean, and shows t h e t i m e o f t h e minimum and maximum d e v i a t i o n s d u r i n g t h e n i t r a t e - d e p l e t e d p e r i o d .  VAB NAME  | |  TEMP 1 TEMPN | N03N 1 OXY J OXYN | SAT 1 CHLA J CHLB I CHLC J CT J BA j CA ] CTA i BC J BCT | CCT | PGO J PNO | RES J PROD j PGODY J PCDY \ PGOST | PNOST 1 RESST | ASS | EXCST | ALPHAG| ALPHACJ  ANALYSIS OF VARIANCE \ BY STATION J BY TIME SIG- j T F SIG. J A F 27 27 27 27 27 27 27 27 27 27 27 27 27 27 27 27 10 10 10 10 10 10 10 10 10 10 5 10 10  MULT 1 GRAND J J MEAN J  MULTIPLE CLASSIFICATION ANALYSIS DEV«N BY STATION J DEV •N BY TIME DAY J MAX. SUB MID BOT OUT J MIN.  *** -000 1 4 3.2 -028 - 02 . 998 J 12-9 | *** -000 1 4 3-2 .029 | . 998 j 2-5 1 .02 *** . 000 1 4 0- 5 -677 J .999 1 19.1 J .00 29. . 000 j 4 7. 2 -000 | .953 | 12.02 i 0.23 -000 .000 .000 - 000 .000 *** -000 20. .00 0 5.6 .000 26. .000 29. .000 31. .000 9.3 -000 25- . 000 18. .000 1.8 - 150 1. 7 . 163 21. .000 1. 4 - 247 9-8 -000 5. 1 . 002 7.7 .000 38. .000 9.9 -003 4.6 -003 9.8 .000 22. 37. 18. 20. 24.  1  1 1 1 1 1 1 1 1 1 1 i 1 I 1 I ] 1 1 1 1 1 1 1 1  4 4 4 4 4 4 4 4 4 4 4 4 3 3 3 3 3 3 3 3 3 3 3 3 3  7-2 -000 i 8.3 -00 0 j 4- 1 -00 9 ! 0.9 .442 | 3.3 .026 j 2.7 .052 | 1-4 -264 2.7 .053 | 1.6 .209 i 1. 4 . 25 2 | 2. 4 -071 | 1.8 - 160 j 2.6 .103 ] 3.0 -076 .23 -786 | .92 .318 J 2. 4 -117 | 0-9 -428 3.8 -022 | 5- 8 .011 ] 0.3 -735 1 1-9 - 176 | 2.6 . 134 10. .00 1 18. .000 i|  .940 -962 . 926 .933 -944 - 876 .933 -815 -947 -953 .956 -870 -96 3 -951 - 688 .697 .957 . 670 .919 . 873 -892 -980 -921 . 880 .934  .01 -.01 .01 -.01 -03 -.01 - 11 -.18 i 4.63 J 0.23 - 11 -. 18 J 135. 1 2-8 1.3 -2.3 1-3 1 40.6 ] -1.8 0-2 .03 . 13 -- 11 j 1-6 I i 20.5 | --24 -.48 .37 0.7 1 45.0 1 -2.0 0-2 .00 1 .04 % .01 -00 -02 -.01 -.01 | -51 | | 1- 12 ) .00 .00 -.02 . 00 1 - 0 8 3 -01 -00 .00 j .04 1 -01 -00 -00 3 .46 | -02 -.01 i 3 4 1 . 1 20- -13. - 7. i 294. | 23- -13. -10. 1 47. 1 ~ 3. - 0. - 4. 2. J 51. J 3- - 4. j 3.17 1 19. -13. -06. - 12 | 0.46 J 25- -37. 1.0 -0.4 -0. 6 1 9.5 | | 8.0 1 1- 1 -0.5 -0.6 | 1.5 | - 0 . 1 0. 1 0.0 0- 1 -0. 1 0-0 1 1-7 \ 0.8 -0.2 -0.6 | 6.0 ] 1 65.4 1-21- 4 -9.5- 30-9 4.4 ] 10.4 1- 3-2 -1-5  24 -.02 | -1.07 1 . 54 31 -.02 | 8 -.02 \ -7. 67 -2.25 23 -.16 j 23 -. 16 J -1.68 23 -1.8 i -26. 9 9 0.3 | -12. 7 -.04 | -1.60 20,21 19 .36 j -7. 23 9 •1.0 | -20. 2 .00 | - .04 19-21 - .09 19,22 .01 9 .02 | - .23 .00 | - .08 20,21 .00 J - . 04 19-21 19 -00. , - . 14 6 -249. N/A 6 -231N/A 9 - 24. N/A 6 N/A : - 16. 6 -2. 26 N/A 6 N/A - . 13 30 - 3.0 N/A - 2. 8 30 N/A - 0.8 24 N/A 0.7 21 N/A | - 2. 8 30 N/A 6 -29.2 N/A 6.4 18 N/A  | 1 | | | | | I | 1 1 1 1 1 1 | I 1 1 1 | I  i  | 1 J 1 3 I  DAY  1-33 13,34 1.04 20 27 3.53 12 1.91 1.71 33,34 13 27.7 11-6 15 3-54 8 24 4.31 9-6 15 8 -11 .09 29,33 .34 26 .21 8 8 - 1 1 .18 9 108. 12 79. 12 29. 12 1834 1.04 12 -13 34 6 4. 6 2. 1 18 3. 0 6 3. 7 6 1.8 18 78.8 30 30 10.5  128  Table  15.  P r o d u c t i v i t y component a n a l y s i s f o r EXE4. RPGC,APGC and EPGO r e p r e s e n t t h e p r o p o r t i o n of g r o s s p r o d u c t i v i t y due t o r e s p i r a t i o n , a s s i m i l a t i o n a n d e x u d a t i o n . , ESTPGO i s t h e e s t i m a t e d g r o s s p r o d u c t i v i t y b a s e d on t h e model: ESTPGO=EPGO * APGC -*• EPGG . See t i e t e x t f o r an e x p l a n a t i o n c f t h e r e s u l t s .  EXP4 - TASK a  V AR  RPGO APGO EPGO ESTPGO RPGO APGO  IG1AND ! MISN j | 0.21 J 0.18 | 0,59 | 0.9 8 I | 0.20 | 0.19  ANALYSIS CF VAEIANCE BY STN EY TIME I TOTAL A SIG T SIG 1 K SIG J 3 .9 88 .055 I 15 .1 19 5- .029 ] 15 .one 3 .271 3 ,002 .002 i 15 .0 01 5 3 .231 .461 | 15 .372  | KDLT ] R , | .805 | .857 | .551 | .697 I j .745 | .816  3 .946 3 .039  10 .048 10 .001  1  I 30 .081 \ 30 .001  EXP4 - TASK E  V AR  RPGO APGO EPGO ESTPGO EPGO APGO  | GRANT; I | MEAN I 1 | 0.18 1 \ 0. 19 1 | 0. €3 1 | 0.99 i  ANALYSIS OF VARIANCE BY STN 1 BYTI ME | TOTAL A SIG I T SIG j K SIG  MOLT R .844 ,953 .768 .762  !  in  j 0.15 \ .807 1 0. 17 I .921  j  1  .387 .893 .526 .872  5 .035 1 5 .000 1 5 .122 1 c .108  3 .434 3 .581  110 .011 I 10 .000  3 3 3 ' 3-  j |  15 .061  15 .o e 1  i  15 .193 15 .2C1  :  30 .016 30 .0 00  129  Table  16,  E x p e r i m e n t a l d e s i g n f o r the i n v e s t i g a t i o n o f t w c stage continuous c u l t u r e s of p l a n k t o n i c herbivorous food c h a i n s , with v a r i a b l e f l u s h i n g r a t e s i n both the primary p r o d u c t i o n tank e x p e r i m e n t s (FXP5A,EXP5B) and t h e s e c o n d a r y p r o d u c t i o n t a c k s (EXP 1 t c E X E 4 ) , R e s u l t s f o r the secondary p r o d u c t i o n systems a r e d i s c u s s e d i n C h a p t e r 7,,  TIME F E S I 0 0 (Days) FBIMABY PEOCDCTION SYSTEMS  t=1, 14 t=15,28 t=29,4 1 t=42,49  EXP  SFCONCABY EECEUCT1CN SYSTEMS  TIME PEBIOD (days)  EXP5A F, £. (/day)  EXP5B F, B, (/day)  0.2 5 0.50 0.10 N/A  1.00 1.00 1.00 0.50  F.R. . (/day)  I  t=18,25  1.00  II  t=26,33  2,00  III  t=35,4 2  1.00  t=43,50  2.00  SOURCE OF INFLOW Tank B (F. E. = 1. 0) Task E (F.B. = 1.0) Tank A (I»B.=0„1) Tank E ( F . B . = 0.5)  PBIBABY COMBO NITY Diatoms Diatoms Flagellate Mixed  130  Tails  17.  R e s u l t s of the n u t r i e n t enrichment e x p e r i m e n t s a t low (0.13 l y / m i n ) and h i g h (0.40 l y / m i n ) i n t e n s i t i e s of p h o t o s y n t h e t i c a l l y a v a i l a b l e r a d i a t i o n during 1XP5. The p r o d u c t i v i t y was measured by t h e u p t a k e o f r a d i o a c t i v e c a r b o n (ug C / l / h r ) . The * V i t a o i n mix' and * V i t a m i n B12* a d d i t i o n s were made a t t h e same concentration (*) o r a t 10x t h e c o n c e n t r a t i o n (**) as i n t h e •! Medium' a d d i t i o n .  EXPEBIMINT 1 (low PfiB) Enrichment  EX IIBIMENT 2 (high PSE)  Productivity  Enrichment  Control  101.  Centre!  E Medium (10.0 mi)  117.  Medium (10.0 ml)  114,  110,  (1.0 ml)  106.  (1.0  wl)  V i t a m i n Mix (+*)  F  93.  Vitamin (**)  10 4.  Mix 103.  (*) V i t a m i n E-12 (**)  112.  Vitanin <**) (*)  Productivity  83. B12 136. 123,  Table  18.  V AR SR SAL TEMP  STN i -  D e s c r i p t i v e s t a t i s t i c s f o r s e l e c t e d v a r i a b l e s f o r EXP5A, i n c l u d i n g a breakdown i n t o t h e t h r e e p e r i o d s o f v a r i a b l e f l u s h i n g r a t e s : F.R.=.25/day f o r t=1,14; F.R.=.50/day f o r t=15,28; a n d F.R.=.10/day f o r t=29,41. N r e p r e s e n t s t h e t o t a l number o f d a t a p o i n t s , i n c o r p o r a t i n g b o t h t h e f a c t o r s TIME and STATION.  |  3 -  |  N  s 41  1 j 2-5 3 TEMPN 8 2-5 I N03 J 1 'i 2-5 3 N03N 2-5 | TN03M | 2-5 | OXY 3 1 1 2-5 J OXYN 2-5 1 TOXIN 3 2-5 J SAT 1 3 2-5 1 CHLA 2-5 1 CHLB 2-5 J CHLC 2-5 J CT 2-5 1 BA 2-5 J CA 2-5 | CTA 2-5 3 BC 2-5 3 BCT 2-5 3 CCT 2-5 |  1  41 41 164 164 41 164 164 164 41 164 164 164 41 164 164 164 164 164 164 164 164 164 164 164  TOTAL EXPERIMENT MEAN S.D. RANGE 404. 27. 2 11. 8 18. 1 6.4 16-7 1.7 14. 9 17.3 7. 24 9.93 2-69 2-95 80. 124. 17.7 0. 5 9.6 26- 0 0. 003 0. 632 1. 779 0. 100 0.040 0.368  105. 6 371. 0.92 3. 1 1.08 4.2 8.5 2.3 8 2-4 5 5.7 5-97 22.0 5.54 21.7 7-68 28.5 27. 1 7.07 ... 787 2. 88 1.629 6.84 1.83 4 7.74 1.849 7.61 10- 0 38. 19-3 8554-0 12-95 0-81 3.6 6-96 39-6 65.8 16.32 0. 157 0 . 833 0.389 2 -667 0.996 6 . 829 0. 148 1 .000 0.059 0-294 0. 123 0 -714  MAX 51928.9 14. 3 21-9 8.4 25- 1 21.7 25. 1 2 7. 1 8. 89 13. 83 7.05 7.05 102. 177. 54. 1 3.6 39.6 66.0 0. 833 2.667 7.800 1.000 0.294 0.714  3 1  F-R-=- 25/DAY MEAN N S. D. j 14  1 1 14 3 56 i 56 3 14 1 56 3 56 1 56 1 14 1 56 J 56 I 56 1 14 i 56 1 56 1 56 3 56 1 56 I 56 3 56 1 56 56 3 56 J 56  495. 12- 0 18. 7 6.7 14.3 5.0 9.3 12.0 7. 65 9.40 1. 74 1. 98 84. 119. 9.7 0. 9 4. 4 12- 6 0.143 0 .586 1.558 0-201 0.085 0.370  14.9 0.76 | 2.16 | 1.55 2.91 J 8.60 J 6. 19 | 6. 12 J -473 1-822 | 1.658 i 1.794 j 6. 1 ] 24.5 1 10-67 ] 1. 10 4-50 | 11-45 J 0.172 | 0.424 j 0. 464 ! 0.189 | 0-075 J 0-179 I  F.R-=-50/DAY MEAN S.D. I N 14 14 56 56 14 56 56 56 14 56 56 56 14 56 56 56 56 56 56 56 56 56 56 56  33 7- .13.1.7 | I 1.49 J 12- 2 1.39 j 16.0 1.26 i 3. 8 7.70 J 14. 1 0.0 0-08 3 14. 1 7-4 8 J 5.72 1 16. 3 7-46 - 973 1 10.64 .813 3 3- 18 1- 53 4 3 3.71 1- 405 J 12.7 J 838.4 j 127. 12.34 J 26.6 0.63 J 0.3 14. 5 7.01 j 34-0 13.72 J 0.017 0.031 | 0. 549 0-08 1 ] 1-322 0.159 | 0.029 0-051 J 0.012 0-020 J 0-419 0-066 j  F.R.=. 10/DAY MEAN S. D. N 13  378.  49.0  13 52 52 13 52 52 52 13 52 52 52 13 52 52 52 52 52 52 52 52 52 52 52  11. 1 19.8 8.8 21.9 0.0 21.9 24. 1 6. 55 9.73 3. 18 3. 18 71. 125. 16.8 0.3 10.0 3 1.8 0- 090 0- 772 2. 511 0. 068 0.021 0.310  0-34 1.61 1.38 1-24 0. 13 1-20 1.98 . 160 1. 812 1. 896 1. 896 2.0 2 0.4 9.52 0. 37 4.87 14. 31 0. 192 0.503 1. 4 43 0- 103 0.033 0.054  Table  19-  VAR SR SAL TEMP  D e s c r i p t i v e s t a t i s t i c s f o r s e l e c t e d v a r i a b l e s f o r EXP5B, i n c l u d i n g a breakdown i n t o t h e two p e r i o d s o f v a r i a b l e f l u s h i n g r a t e s : F-E-=1.00/day f o r t=1,41 a n d F.R-=-50/day f o r t = 4 2 , 4 9 . Note t h a t t h e f i r s t p e r i o d (F. R«=1. 00/day) c o r r e s p o n d s t o t h e t o t a l e x p e r i m e n t a l p e r i o d f o r EXP5A. N r e p r e s e n t s t h e t o t a l number o f d a t a p o i n t s , i n c o r p o r a t i n g b o t h t h e f a c t o r s TIME a n d STATION.  1 STN J  1  i i i TEMPN i N03 i i N03N i TN03N i OXY i i OXYN TOXYN 1 SAT 1 1 CHLA 1 CH LB a CHLC 5 CT i BA i CA ? CTA i BC i BCT i CCT i  ]  N  j  41 41 41 164 164 41 164 164 164 41 164 164 164 41 164 164 164 164 164 164 164 164 164 164 164  1 1 12-5 2-5 1 2-5 2-5 2-5  1 ] | J | | | j  2-5 2-5 2-5 1 2-5 2-5 2-5 2-5 2-5 2-5 2-5 2-5 2-5 2-5 2-5  i ] J J | 1 1 | | J | J | \ ]  F-R. = 1.00 PEE DAI MAX MEAN S. D. RANGE 404. 27-2 11.9 15.0 3-1 16.4 2.8 13.5 15. 9 7.29 10-92 3.62 3.87 80128. 25-8 0.6 15-0 30.0 0. 081 0. 659 1.275 0.093 0. 049 0.500  105-6 0.92 1-10 .925 0.87 6. 18 6.48 8.92 8-06 .884 1.612 1-999 1.882 11. 1 19. 4 15.78 0.88 9.76 17.67 0- 172 0.392 0.396 0- 133 0.075 0. 156  371. 3- 1 4.2 3.8  4-3  2 2. 4 21-9 29.8 27. 1 3.08 6-44 8.08 7-49 38. 74. 61.4 3.3 39. 1 6 9.3 0.9 43 2- 285 2-359 0-545 0-318 1.013  519. 28.9 14. 3 16. 5 4. 9 24.9 21.9 24. 9 27. 2 8.96 14. 12 7.89 7.89 101. 88. 61-5 3-3 39- 1 69.5 0.943 2-28 5 3.20 1 0.545 0.318 1-Q13  1  N  I i  8 8 8 32 32 8 32 32 32 8 32 32 32 8 32 32 32 32 32 32 32 32 32 32 32  1  a i i  i  \  1 1 1  I I 1 J 1  1 ] 1 9 8 1 1 1  F.E-=0-50 PEB DAY MAX MEAN S.D. RANGE 298.  89.0  259.  380-  10.6 15.0 4. 4 22.9 0.0 22. 9 23.4 6. 14 11.05 4-90 4-90 66. 13032.7 0.7 20. 1 38.3 0-023 0-616 1.169 0.036 0-020 0.528  0. 46 0.57 0.83 1.86 0.0 1.76 1.89 0.387 0.981 0- 908 0.908 4.62 1 1.6 3-96 1-52 2- 95 5. 14 0- 049 0. 084 0. 073 0. 077 0. 041 0.076  1.4 1.8 2.5 5-0 0.0 5.0 5.6 1-00 4-44 3. 86 3. 86 12. 51. 15.9 7.0 14.5 22. 5 0-227 0-418 0.341 0.370 0.189 0.379  11.5 16. 0 5.6 2 5.4 0.0 25.4 26.6 6.76 12.61 6.27 6.27 74. 148. 41.0 7.0 30. 3 48.0 0.227 0. 959 1.338 0- 370 0-189 0-828  Table  20.  R e s u l t s o f t h e a n a l y s i s o f v a r i a n c e and m u l t i p l e c l a s s i f i c a t i o n a n a l y s i s f o r s e l e c t e d v a r i a b l e s a s a f u n c t i o n o f t h e i n d e p e n d e n t f a c t o r s TIME a n d STN f o r EXP5A. S t a t i s t i c s a r e b a s e d on d a t a f r o m t h e n i t r a t e - d e p l e t e d p e r i o d (T>4). *** i n d i c a t e s F - v a l u e s g r e a t e r t h a n 99. The e l e v e n 'TIMES• f o r t h e p r o d u c t i v i t y v a r i a b l e s i n c l u d e e v e r y t h i r d day from Day 6, e x c e p t Day 15 when t h e d a t a was m i s s i n g . The MCA i n d i c a t e s t h e e f f e c t o f each c a t e g o r y o f STATION, e x p r e s s e d as a d e v i a t i o n from t h e g r a n d mean, a n d shows t h e maximum and minimum deviations d u r i n g t h e n i t r a t e - d e p l e t e d p e r i o d . MULT R i s t h e m u l t i p l e c o r r e l a t i o n between t h e d e p e n d e n t v a r i a b l e and b o t h i n d e p e n d e n t v a r i a b l e s TIME and STN. S i g n i f i c a n c e v a l u e s i n t h e A NOVA a r e b a s e d on t - 1 d f f o r TIME and a-1 d f f o r STATION.  VAfi NAME  J ]  TEMP J TEMPN ] N03 1 N03N 1 TN03N 1 OXY i OXYN J TOXYN | SAT 1 CHLA 1 CHLB 1 CHLC i CT | BA j CA J CTA 1 BC 1 BCT 1 CCT | PGO J PNO | EES \ PROD 1 PGODY | PCDY I PGOST | PNOST | RESST | ASS 1 EXCST | ALPHAG| ALPHACJ EPGO | APGO |  ANALYSIS OF VARIANCE J BY STATION | BY TIME SIG. SIG. 1 T F F 1 A 37 37 37 37 37 37 37 37 37 37 37 37 37 37 37 37 37 37 37 11 11 11 11 11 11 11 11 11 11 5 11 11 11 11  MULT | GRAND J MEAN H  |  j  MULTIPLE CLASSIFICATION ANALYSIS | DEV • N BY TIME DEV* N BY STATION MIN. J MAX. SUR BOT OUT j DAY MID  *** -000 ] 4 41. .000 i .994 1 18. 4 0. 15 0.09 -.45 0.20 41. -000 | .995 *** .000 J 6.6 | 0. 15 0. 10 -.45 0.20 ! ^  2.5 .06 1 *** -000 *** .000 *** .000 *** .000 *** .000 *** .000 *** .000 7-3 .000 43. .000 90- .000 8.6 .000 6- 1 .000 26. .000 5.2 -000 6.3 .000 4.0 -000 18. .000 18. .000 1.7 . 153 34. -000 14. . 000 26. .000 3-4 .010 12. .000 4.3 .003 3.7 .006 8.3 .006 2-4 .043 2. 6 -032 7-1 .000 3-2 .012  i 4 1.0 . 482 | 4 2. 8 .042 1 4 2-7 .051 1 4 12. .000 1 4 12. .000 1 4 12. .000 1 4 18. .000 1 4 4.9 -003 ! 4 1- 2 .31 1 ! 4 3.2 .025 i 4 2.3 .078 ] 4 .79 -500 t 4 1. 5 -213 i 4 .62 .606 1 4 .42 -737 I 4 .36 .782 1 4 3.0 .033 1 3 1.3 -284 1 3 0.7 -523 1 3 1.4 .26 7 1 3 1.7 .208 i 3 1.4 -261 \ 3 1.8 . 197 1 3 1. 1 -35 3 1 3 1-4 . 267 1 3 1- 4 .259 I 3 0. 5 .626 1 3 1. 8 . 234 | 3 0.2 .813 1 3 3. 1 .068 ] 3 1.9 . 170 1 3 1-5 .23 8  J .536 1 0.0 J1.000 I 16.5 3 1.000 I 18.8 3 - 9 9 1 3 10.05 J -993 J 2.83 j .993 3 3.09 J .987 3 126. I .99 1 3 18.8 1 -851 3 0.4 i -969 3 10.2 i - 9 8 5 1 27.9 | .862 3 0.07 j . 829 3 0.61 3 -951 J 1.81 ] .798 3 0.08 3 -824 1 0.03 3 -769 J 0-35 J -949 3 1601 .948 3 119. 41. 3 -705 | 1 - 972 | 61. 3 - 938 | 1.39 | . 983 | 0.50 J .80 1 I 1 0 - 6 | .929 J 6.5 J .834 J 4. 1 3.8 3-811 1 J - 906 1 2.2 | .744 3 39.8 1 .786 3 14.0 1 .888 1 0.32 1 .799 ] 0.40  3 1 j j j  ] 1 1 j 1 1 3 j I j 3 1 j j 1 3 j j 3  J i  i  -.01 0.01 0.01 0- 13 0. 13 0-13 2.0 -0.6 0.0 -0.3 -0,6 0.01 0. 04 0.01 -.01 0.00 0.01 4. -3. 6.6 -2. 4 .04 -.25 2- 1 -0. 1 2.2 0- 3 -0-2 -1.6 -2. 5 0. 06 -.04  -.01 .03 0.01 -.03 0.01 -.04 0-04 -.21 0-04 -.21 0. 04 -.21 0.8 -3.6 -0.5 -1.0 0.1 0.0 -0.2 0.9 -0.8 0-5 0.01 0.00 0.02 0-02 0.05 - . 06 0.01 0-00 0.00 0.00 0.00 0.03 9. - 1 3 . 9. - 6 . 0.2 -6.8 4.2 -1.7 .08 - . 12 .34 -.09 -0.4 -1.8 0 . 6 -0.5 -0. 9 -1.2 0.0 -0.3 0.3 - 0 . 1 2-9 -1.3 2-7 -0.2 -.03 -.03 -.02 .05  -.01 j 0.0 1 0.01 0.04 | 0.04 0.04 0.9 , 0. 1 , -0. 1 | -0.4 | 0-9 -.02 \ -.07 J 0.00 j 0.00 | 0.00 | -.0 3 J N/A j N/A ; N/A ;  N/A N/A N/A N/A N/A N/A  | ! : J i j J :  N/A N/A N/A N/A N/A i N/A ,  -4- 5 27 | 20 J -5. 1 0-0 MANY J 20 J -13. 4 -9- 2 20 | 41 1 -3.02 -3.35 10 | -3.47 10 3 -34. 41 j -18.0 41 | -0. 4 23-31 3 -9.5 11 J -23. 5 41 | -0.07 MANY J -0. 47 1 1 3 -0. 68 5 | -0. 08 23-31 1 -0.03 23-31 3 -0. 26 11 | 9 | -104. -107. 9 J 39 | -20. 9 9 J -46.9 -- 89 9,39 3 9 3 38 -6. 0 18 a -3.6 18 1 24 | -2. 8 18 ] -2. 1 -0.7 18 1 -17. 9 36 3 -6.4 27 J -.22 21 3 -. 18 21 J  2.9 3.6 0.2 8.6 8-4 3.58 3.91 3- 65 48. 26.8 3-0 16-6 28.7 0.55 1.58 4.30 0. 41 0. 11 0. 14 109. 125. 21-4 50- 1 1- 03 .36 1 7 . 4  7.2 18.5 3.0 1-3 29-2 8.9 0.47 .29  DAY 37 36 33 27 34 5 29 29 5 26 5 26 26 41 41 41 8 9 24 21 21 27 24 21 6 9 21 9 9 12 6 24 9 6  T a b l e 21.  R e s u l t s o f t h e a n a l y s i s o f v a r i a n c e and m u l t i p l e c l a s s i f i c a t i o n a n a l y s i s f o r s e l e c t e d v a r i a b l e s a s a f u n c t i o n o f t h e i n d e p e n d e n t f a c t o r s TIME and STN f o r EXP5B. S t a t i s t i c s a r e b a s e d on d a t a from t h e n i t r a t e - d e p l e t e d p e r i o d (T>6). F - v a l u e s g r e a t e r t h a n 99. a r e d e n o t e d by The e l e v e n •TIMES' f o r t h e p r o d u c t i v i t y v a r i a b l e s i n c l u d e e v e r y t h i r d day f r o m Day 6, e x c e p t Day 15 when t h e d a t a was m i s s i n g . The MCA i n d i c a t e s t h e e f f e c t o f e a c h c a t e g o r y o f STATION, e x p r e s s e d a s a d e v i a t i o n from t h e g r a n d mean, and shows t h e maximum and minimum deviations d u r i n g t h e n i t r a t e - d e p l e t e d p e r i o d . MOLT R i s t h e m u l t i p l e c o r r e l a t i o n between t h e d e p e n d e n t v a r i a b l e and b o t h i n d e p e n d e n t v a r i a b l e s TIME and STN. S i g n i f i c a n c e v a l u e s i n t h e ANOVA a r e b a s e d on t - 1 d f f o r TIME a n d a-1 d f f o r STATION.  VAR NAME  J ]  TEMP ] TEMPN | N03 ] N03N J TN03N I OXY J OXYN | TOXYN J SAT J CHLA | CHLB J CHLC ] CT J BA J CA | CTA J BC | BCT | CCT J PGO J PNO j RES | PROD J PGODY | PCDY 1 PGOST | PNOST | RESST J ASS | EXCST J ALPHAG] ALPHAC]  ANALYSIS BY T I M E T F SIG. 35 35 35 35 35 35 35 35 35 35 35 35 35 35 35 35 35 35 35 11 11 11 11 11 11 11 11 11 11 5 11 11  OF VARIANCE | BY S T A T I O N MOLT H S I G . ,| 1 A 7  *** . 0 0 0 *** . 0 0 0 37.  .000  *** . 0 0 0 *** - 0 0 0 16. 30.  *** 1550. 47. 68. 42. 21. 24. 21. 19. 17. 16. 83. 653.2 9.6 68. 8.9 15212. 7 8.8 598.0 3. 1  .000 -000 .000 .000 .000 .000 -000 .000 -000 -000 .000 -000 .000 .000 .000 .000 -012 -000 -000 -000 .000 .000 .029 .000 .000 .000 .015  |  1  )  {  j j j 1  | ] | |  4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 3 3 3 3 3 3 3 3 3 3 3 3 3  25. 251. 7 1.7 1. 7 11. 11. 11. 12. 3.7 - 38 1- 2 1- 1 -28 .98 1.3 .41 .35 . 57 0. 8 0. 4 1.6 0. 5 0. 8 0. 5 2- 1 0. 7 0.2 1-0 0. 1 37. 22.  .000 -000 .168 -163 .164 -000 .000 -000 -000 .015 .768 .304 .35 2 -843 -403 .265 -748 -789 -637 -46 5 .666 -235 -607 .473 -610 .151 -507 -842 .403 -939 .000 .000  ; ] j | | j ] I J |  | j | | | | | | ; J | ; !  -986 .986 .962 . 999 .999 .921 .955 .947 .920 .971 .970 .979 .966 -934 .943 .935 -928 .922 .919 -988 -985 -800 -911 .986 .905 . 941 .956 .759 .904 -984 .941 .888  j GRAND | | MEAN J  MULTIPLE CLASSIFICATION ANALYSIS D E V N BY T I M E DEV•N BY S T A T I O N I SUR OUT I MIN. | MAX. MID BOT DAY  1 1 5 . 1 J 0. 1 2 I 3.2 | 0 . 1 2 | 0.3 1 0.0 0-0 | 16.2 | J 0 .0 j 18.3 111-38 1 0 . 2 4 | 4.17 1 0.24 J 4 . 4 1 1 0.24 I 134. ] 3.2 | 2 9 . 6 I -0.6 ] 0.6 1 0-0 0.0 1 17-1 3 0.0 i 34-3 1 J 0.03 1 0.00 | 0.57 ) 0.01 | 1. 1 8 I 0 - 0 2 1 0.06 1 0.00 1 0.02 1 0-00 3 0.48 1 0-00 1 3 0 4 . ,1 7. 1 263. 1 -4. | 44. | 8.5 1 101. J " 5 .06 | 2. 56 1 I 0 - 8 4 | -- 4 6 | 11-0 1 0-2 0.0 1 9.5 | 0-0 1 1-7 I | 3.8 I - 0 . 4 i 4.0 ] 0.0 1 48.1 1-15.4 1 15.7 1 - 6 . 3  -.02 -.02 0.0 0.0 0.0 0.21 0. 2 1 0-21 2.4 -0.4 0,0 0. 1 -0.5 0.00 0.01 -.01 0.00 0.00 0.01 0. -1. -2. 1 -3.00 -.23 0-6 0.3 0- 1 0.2 -0. 1 -1.7 -0.6  -.21 -.21 0- 1 -0.1 -0. 1 -. 44 -. 4 4 -. 44 -5.7 1.8 0.0 0-4 1-1 0.00 -.01 -.01 0.00 0-00 0.00 -7. 5. -6-4 8. -.06 .70 -0.8 -0.3 -0. 1 0-2 0. 1 17.1 7-0  0-10 0- 1 0 0.0 0.0 0-0 -.01 -.01 -.0 1 0.2 -0.9 0.0 -0.5 -0.7 0.00 0.0 0-01 -.01 0-00 -.01 N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A  j -2. 1 ! -2.6 | -0. 3 i -13. 7 | -9. 3 | -2. 80 | -2. 84 -2. 92 | -32. 2 3. 3 | 0.6 ] | -15. 0 | -26.8 | -0-03 | -0. 34 | -0. 25 J -0. 06 \ -0. 02 J -0. 25 ; -225. | -200. -37. 2 -64. i i - 1.85 i -. 5 2 ; -5.9 -5. 3 | j -1.3 ; -2- 0 j -2. 6 | -19.7 -8.2 |  26 20 MANY 20 20 12 20 12 12 12 MANY 11 12 MANY 11 7 MANY MANY 1 1 12 12 6 12 12 12 18 18 6 27 12 21 21  1 J | 1 1 | | J 1 i 1 | 1 1 1 1 J \ | J 1 | J 1 1 1 1 1 | | 1 I  1. 3 1.4 4.0 8.5 8.9 1.95 2.96 2.72 25. 24. 6 2.3 15.2 26-8 0-07 0.31 0.36 0.27 0.08 0. 1 9 155. 148. 25.2 139. 1.27 1-12 6- 1 4.9 1- 4 4-0 4.2 17. 9 6.2  DAY 9 41 13 27 34 32 32 32 8 27 7 26 27 8 21 36 8 8 18 33 33 36 27 33 27 36 36 21 6 36 30 9  137  Table  22.  R e s u l t s f o r the p r o d u c t i v i t y v a r i a b l e s , averaged f o r t h e t h r e e p e r i o d s o f v a r i a b l e f l u s h i n g r a t e s i n .EXP5A, w i t h EXP5B r e s u l t s d u r i n g t h e sane t i m e p e r i o d as a c o m p a r i s o n . S a m p l i n g t i m e s i n c l u d e Days 6 , 9 and 12 f o r P e r i o d 1 ( F . I . = . 2 5 / d a y ) , Days 18,21,24,27 f o r E e r i o d 2 (F.B.=,50/day) and Days 30,33,36,38 f o r P e r i o d 3 (F.B. = . 10/day) . Means f o r t h e t o t a l t i m e p e r i o d a r e f o u n d i n T a b l e 20 and T a b l e 21,,  EXP5A VAB  1 1 PGO 1 PGO 1 EES 1 FBOE I PGOST 1 PNOST 1 E E S ST 1 ASS 1 PGODY 1 FCDY. 1 ALP BAG| ALPHAC! APGO J EPGC 1  STN 1  F.B.=.25/LAY N MEAN S.D.  2-4 2-4 2-4 2-4 2-4 2-4 2-4 2-4 2-4 2-4 2-4 2-4 2-4 2-4  9 9 9 9 9 9 9 9 9 9 9 9 9 9  J | | | | | i | J j J | { |  F.E.=.50/DAT N MEAN S. D. ;1  |  111. 46.3 ! 12 69. 45. 1 1 12 4 2. 20.6 I 12 56. 40. 2 12 16.6 13.28 12 7. 1 3.06 I 12 9.5 13. 1 | 12 6.0 2.3 12 1.00 - .414 | 12 0.47 .322 I 12 43.3 31.39 J 12 15.3 5.68 | 1 2 0.45 .203 | 12 0. 4 5 . 2 7 6 1 12  224. 182. 42. 70. 8.4 7.0 1.5 2.5 1.96 €.58 47.7 15.4 0.34 0.21  88.4 85.8 21.3 30.6 4.40 4. 4 2 0.70 0.94 .592 .173 17.72 8,61 .145 .146  F,I.=.10/tA 1 MEAN •£.C. . M •12 12 12 12 12 12 12 12 12 12 12 12 12 12  132. S3. 40. 55. 8.2 5.5 2.6 3.3 1. 1 1 0.45 29.3 11.7 0.43 0.32  6 2.0 54.9 22. 1 2 3.6 2.25 1.79 1.30 0.70 .5 1 3 . 1S5 1 1. 1 2 3.45 - 135 .142  I 12 I 12 12 12 j1 1 2 | 12 12 12 12 | 12 I 12 12 12 12  420. 366. 55. 91. 13.7 11.9 1.8 2.9 3.50 0.75 57.7 12.7 0.22 0.13  56.1 14.2 22.4 26.3 2.77 2.09 0.66 0.5 2 .4 6 7 .218 16.56 6.22 .064 .045  I  | I |  i  EXP5E PGO 1 PGO J EES 1 PBOD 1 PGGST 1 PNOST 1 EESST 1 ASS 1 PGOPY 1 PCDY 1 ALPHAG| A L P B AC| AIGC 1 EPGO 1  2-4 | 2-4 | 2-4 | 2-4 J 2-4 | 2-4 | 2-4 | 2-4 I 2-4 J 2-4 | 2-4 | 2-4 | 2-4 | 2-4 )  9 9 9 9 9 9 9 9 9 9 9 9 9 9  2 0 6 . 1G5.6 | 179. 89.9 | 27. 26.8 | 110. , 68.7 | 1.7 0 I 12.6 10.9 1.91 1.7 1.11 I 1.90 | 6.5 1.78 .913 I .597 0.93 35.5 10.68 | 9.67 | 18.7 0.52 .047 | 0. 1 3 . 1 0 6 |  12 12 12 12 12 12 12 12 12 12 12 12 12 12  262. 134.2 2 2 5 . 116.8 46. 26,6 105. 88.2 7.1 1.8S 6. 1 1.74 1.07 1.5 2.6 1. 1 2 .874 2.20 0.87 .704 46. 1 24.42 16.4 8. 2 6 0.38 .156 0.20 . 105  138  Table  23.,  P r o d u c t i v i t y component a n a l y s i s f o r IXP5. RPGO,APGO and EPGO r e p r e s e n t t h e p i o p c r t i o n o f g r o s s p r o d u c t i v i t y due t o r e s p i r a t i o n , a s s i m i l a t i o n and e x u d a t i c n . ESTPGO i s t h e e s t i m a t e d g r o s s p r o d u c t i v i t y b a s e d on t i e s u b - m o d e l : ESTPGO=BPGO • APGO * EPGO . See the t e x t f o r an e x p l a n a t i o n o f t h e r e s u l t s .  EXP5 - TASK A  |GRAND 1 MIAN  BUIT R  ANALYSIS OF VARIANCE BY STN EY TIME TOTAL A SIG T SIG N SIG  | j | |  0.29 0.48 0.29 1.06  .798 .791 .657 ,825  3 3 3 3  RPGO ] 0.32 APGO | 0.40  .888 .799  3 . 170 3 .238  VAR RPGO APGO EEGO ESTPGO  .062 .4 89 .54 9 .724  5 5 5 5  .287 .089 .363 .042  11 .000 11 .012  15 15 15 15  .13 1 .145 .475 .0 86  3 3 .0 0 0 33.016  EXP5 - TASK B  VAR RPGO APGO EPGO ESTPGO RPGO APGO  JGRAND l MEAN  MULT 1  ANALYSIS OF VARIANCE BY STN J BY TIME TCTAI A SIG | T SIG N SIG  | J | I  0.14 0.36 0.34 0.8 4  .856 .951 .585 .781  3 3 3 3  ] 0. 1 5 J 0.36  .764 .890  3 .986 3 .4 27  I  .063 .352 .081 .353  .063 .000 .000 .122  15.046 15 .000 15 .000 15 .165  11 .024 11 .000  33 .0 45 33 .000  5 5 5 5  Table  24.  T w o - s t a g e c o n t i n u o u s c u l t u r e o f h e r b i v o r e s . S t a t i s t i c a l summary o f e n v i r o n m e n t a l v a r i a b l e s d u r i n g e a c h o y s t e r e x p e r i m e n t (t=8) f o r t h e f o u r t a n k s (Tank 4= C o n t r o l ) . ANOVA's a r e f o r Tanks 1,2 and 3 o n l y . See A p p e n d i c e s 1 a n d 2 f o r a d e s c r i p t i o n o f t h e v a r i a b l e s .  EXP. VAR TEMP  STKUP  CTA  EXP.  II  EXP.  III  J  EXP.  | IV  i  ANOVA BY EXP TANK  S.D.  |  S1G. SIG.  |  .000 .793  |  .000  .050  |  .000  . 186  J  .000  .055  |  .000  .970  MEAN  S.D.  MEAN  S.D. |  MEAN  18.4 18. 5 18. 4 18. 4  2.44 2.40 2.36 2.38  | | | |  20. 1 20. 1 20.4 20.0  1. 1. 1. 1.  17 12 16 16  23.3 23.3 23.5 23.1  2.29 2.31 1.86 1.96  J | l |  19.5 19.4 19.8 19. 4  1.51 1.53 1.67 1.48  21.2 22. 6 23. 1 15.3  14.79 14.88 15.47 10.40  | J J J  17.7 25.4 30.4 1 2. 1  8.80 11.60 14. 49 7.00  7.2 8.1 8. 1 5.8  6.25 | 6.86 1 6.92 i 4.9 6 |  18.5 26. 2 29.7 10.3  6.7 1 7.69 9.60 5.73  . 195 .269 3 | .326 4 1 .070  .1947 .1306 .1569 .0833  | | | |  . . . .  031 062 056 000  .0468 .0744 .0579 .0000  1  1.375 1.180 1.258 1.276  | J ] |  4.47 5 . 57 6.31 4.37  1.198 1.570 1.699 1.065  . 120 .326 .285 .256  1 | | I  1.50 1.55 1.47 1.54  1  1  2 1 3 I  4  ]  1  J  4  OXYUP  |  S.D. j  TANK | MEAN  2 I 3 I  BA  I  I  1  ] 2 I  J  3. 39  2 1 3 . 84 3 | 4 . 07 4  1  1.87  1  |  1.35 1.49 1. 55 1.56  2 I 3 J 4  ]  . . . .  400 668 463 229  .4416 .4514 .3044 .2440  | J J J  .019 .068 . 157 .014  .0202 .0557 .1274 .0223  , | |  3 . 97 4 . 16 3.43 3.74  1.112 1.662 1.231 0.898  | | j |  4.77 5- 18 5 . 74 4.24  0.793 0.664 0.566 0.757  .219 | . 199 | .220 ] .181 J  3-80 3 . 72 3 . 76 4.92  2.461 1.038 0.755 2.060  J | I |  1. 48 1.45 1. 57 1.52  . 056 .091 .188 . 152  T a b l e 25.  D e s c r i p t i v e s t a t i s t i c s f o r t h e s i x growth v a r i a b l e s f o r e a c h c u l t c h a t t h e s t a r t o f e a c h o f t h e f o u r h e r b i v o r e e x p e r i m e n t s ; TIME=4 summarizes t h e f i n a l m e a s u r e m e n t s . WIDTH S. D. MEAN  TOTAL WGT DEPTH MEAN MEAN S.D. S.D. •..  MEAT WGT MEAN S.D.  SHELL WGT MEAN S-D.  I.D.  LENGTH MEAN S.D.  TIHE=0 CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH  ALL 1 2 3 4 5 6 7 8 9 10 11 12  5.7 4.0 5. 1 5. 1 4.9 5.2 6.0 5.8 5.8 5.8 6.9 7.2 7. 1  0.97 0.39 0.27 0.40 0. 19 0.39 0. 19 0.26 0.43 0. 16 0.36 0.26 0.37  3.2 2-8 3-4 2.8 3. 1 2-9 3- 6 3.2 3-1 3- « 3.9 3.6 3-2  0. 61 0- 40 0- 35 0- 62 0. 53 0- 50 0. 34 0- 45 0- 83 0- 42 0. 73 0. 70 0. 39  1.7 1-4 1.7 1.7 1.5 1.7 1.6 1.7 1.9 1.6 1.9 1.9 1.9  0-30 0-21 0-15 0-24 0.20 0-36 0.31 0-23 0.24 0.31 0.31 0.29 0.24  12.8 5.7 10.5 9.7 8.9 10.2 14. 4 12.8 14. 1 13.2 18.3 18.3 18. 1  4.70 0.97 1-66 3.59 2-01 2-38 2- 88 2.37 3.50 2.85 3.8 4 2-29 3.60  4.0 1.8 3.6 3- 1 2.8 3-2 4.7 4.0 4.3 4. 1 5.6 5.5 5.4  1.51 0. 38 0.76 1.26 0.9 1 0.75 1. 10 0.94 1.16 1-05 1. 41 1.17 1.20  8. 8 3.9 7. 0 6.6 6. 1 6. 9 9.7 8- 8 9.8 9. 1 12.7 12.8 12.7  3-27 0.68 0.98 2.37 1. 18 1.72 1.88 1. 52 2-48 1-80 2.46 1-50 2-55  96 8 8 8 8 8 8 8 8 8 8 8 8  TIME=1 CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH  ALL 1 2 3 4 5 6 7 8 9 10 11 12  5.8 4.1 5.3 5. 1 5.0 5-3 6.1 6-0 5.8 6.0 6.9 7.3 7.1  0.95 0-38 0.35 0.39 0-21 0.42 0.3 0 0.29 0.39 0.20 0.37 0-29 0.42  3.4 2.9 3.5 2.9 3. 1 3-0 3-8 3. 5 3-2 3-5 3. 9 3.7 3.4  0. 61 0. 40 0. 55 0. 59 0. 47 0. 44 0- 31 0. 53 0. 84 0. 39 0. 74 0. 67 0. 37  1-8 1.5 1-8 1-7 1.6 1-7 1.7 1.8 1.9 1-7 2.0 1-9 1.9  0.28 0. 18 0.12 0. 24 0. 19 0-36 0.26 0-23 0-23 0-28 0.28 0.26 0.23  14.1 6.7 12-0 10-6 10-0 1 1-2 15-7 14.4 15. 1 15.2 19.7 19.7 19. 1  4-87 1- 1 1 1.74 3-77 2.06 2-37 2-74 2.70 3-61 2-87 4. 1 1 2-64 3-99  1.56 4.5 0.41 2. 1 4. 1 0.73 1.29 3-6 0-87 3.3 0.80 3-5 1.05 5.2 0-98 4.6 4.6 1.22 4.9 ' 1.01 1-49 6. 1 6- 1 1. 25 1-28 5.9  9.6 4.6 8.0 7. 1 6.8 7. 6 •10. 5 9.8 10. 4 10. 3 13. 7 13-6 13- 3  3-37 0.78 1.06 2- 45 1-26 1-81 1.80 1.78 2.52 1. 89 2-66 1.68 2.82  96 8 8 8 8 8 8 8 8 8 8 8 3  N  o  TIME=2 CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH TIME=3 CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH  WIDTH DEPTH TOTAL WGT MEAN S- D. ,MEAN S-D. S . D. , MEAN  I.D.  LENGTH MEAN S. D.  ALL 1 2 3 4 5 6 7 8 9 10 11 12  6. 0 4.4 5.6 5.3 5.3 5.4 6.4 6.2 5.9 6. 1 6.9 7.4 7.2  0.93 0.5 0 0-52 0.38 0.37 0-47 0-45 0.33 0.40 0.28 0.36 0.35 0.50  3. 6 3- 2 3.9 3.0 3.3 3. 1 4. 1 3.8 3.4 3. 8 4. 1 3.9 3.7  0.67 0. 55 0.62 0.54 0.41 0. 40 0.38 0-69 0.90 0.50 0.72 0.74 0-45  1.8 1.6 1-9 1.7 1.6 1-8 1.8 1.9 1.9 1-9 2.0 2-0 2.0  0.25 0. 16 0- 12 0. 18 0-23 0.33 0-22 0.24 0-24 0.24 0.26 0.25 0-21  15.9 8.1 14.3 1 1.7 12. 1 12.3 17. 4 16.2 16.7 16.9 22.6 21-6 21. 1  5. 27 1.36 2. 10 4. 1 1 2.04 2.4 8 2.76 3- 18 3-98 3.02 4-72 3. 19 4.75  5.1 2.6 4.8 4-0 4. 1 3.9 5-9 5-2 5.2 5.5 6.9 6.8 6-7  ALL 1 2 3 4 5 6 7 8 9 10  6.1 4.5 5-6 5.5 5.3 5-4 6.4 6-2 6- 1 6-2 7- 1 7.3 7.3  0.93 0.45 0.51 0.43 0.34 0.51 0.4 3 0.33 0.46 0.38 0-28 0.37 0.48  3.8 3. 6 4. 1 3.3 3. 3 3.2 4-2 3-9 3-6 4-0 4.3 3.9 3.9  0.71 0.61 0-70 0.59 0-40 0-48 0.51 0-72 1.03 0.60 0.77 0-74 0.50  1.9 1.6 2-0 1.7 1.7 1.9 1-8 1.9 2-0 1-9 2- 1 2.0 2-0  0-27 0-21 0.23 0.20 0.21 0-36 0.21 0.20 0. 18 0. 33 0.26 0-26 0.22  17.2 9.3 15-5 12.7 12.7 13-2 18.4 17-4 18.0 1 8- 7 24.8 22-9 22- 1  5-66 1.53 2.40 4. 19 2-87 2-75 3- 17 3. 46 4.37 3.37 5.09 3-65 5-13  5-4 2-8 5.2 4.2 4. 1 4.0 5.8 5.4 5.4 6.0 7-4 7. 1 6-9  11  12  MEAT WGT •• MEAN S.D.  SHELL WGT MEAN S-D.  N  1.71 0. 46 0.69 1.57 0.83 0.86 1.03 1.06 1.39 1-03 1.69 1-36 1.49  10.8 5. 6 9.4 7.7 8. 1 8. 4 11. 5 11.0 11. 5 11. 4 15. 6 14. 7 14. 4  3. 63 0.97 1.48 2-58 1- 35 1. 73 1.90 2. 15 2-70 2-05 3.08 2.04 3-36  96 8 8 8 8 8 8 8 8 8 8 8 8  1.78 0.58 0.82 1.53 1-11 0.95 1-23 1. 11 1. 40 1.08 1.60 1. 44 1.51  11. 8 6.5 10. 3 8. 6 8.6 9. 3 12- 6 12. 0 12. 6 12.7 17.4 15.9 15. 3  3. 96 1-16 1.70 2.69 1.85 1.94 2. 11 2-38 3.04 2-35 3.57 2.40 3.73  96 8 8 8 8 8 8 8 8 8 8 8 8  TIME=4 CULTCH, CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH CULTCH  I.D.  LENGTH MEAN S.D.  ALL 1 2 3 U 5 6 7 8 9 10 11 12  6. 1 4.6 5.6 5.5 5.3 5.4 6-5 6.2 6-1 6.3 7- 1 7.3 7-4  0-91 0.4 3 0-4 1 0.41 0.31 0.43 0-41 0-33 0.49 0-35 0-23 0-39 0.48  WIDTH MEAN S . D. ,  3.8 3-7 4. 1 3-2 3-3 3- 2 4-3 3.9 3-6 4-0 4.3 3.9 4-0  0- 74 0. 71 0. 64 0. 85 0. 41 0. 46 0. 47 0. 69 1. 04 0. 51 0. 79 0. 76 0. 45  T O T A L WGT DEPTH MEAN S- D. . M E A N S . D-  1.9 1.7 2.0 1.8 1.7 1.8 1.9 1-9 2. 1 1-9 2. 1 2- 1 2.0  0.27 0.21 0.22 0.20 0.25 0. 33 0.22 0.21 0.20 0.32 0.22 0.23 0.23  19.2 10.5 18. 1 14.5 14.6 14.6 20.8 19.7 19.5 21.6 27.0 25.2 24.4  6. 15 1.77 2.9 4 4.51 2-96 3-19 3-21 4.29 4.97 3-58 5-42 4-50 5-96  M E A T WGT S.D. MEAN  6,1 3.2 5.8 4.9 4.7 4.5 6-7 6.3 6. 1 6-7 8-4 7-9 7.7  1.97 0- 57 0-7 2 1.69 1- 24 1. 13 1-21 1-34 1-57 0-98 1.73 1-64 1.75  SHELL MEAN  WGT S.D.  N  13.2 7. 4 12. 3 9.7 9.9 10.1 14. 1 13. 4 13.4 14.8 18. 5 17. 3 16.7  4.27 1.30 2-44 2-85 1.78 2.23 2-26 3.00 3.46 2.81 3-77 2-99 4.32  96 8 8 8 8 8 8 8 8 8 8 8 8  NJ  143 Table  26. R e s u l t s c f t h e n e t and p e r c e n t i n c r e a s e s p e r o y s t e r f o r t i e s i x growth v a r i a b l e s , i n c l u d i n g s i g n i f i c a n c e l e v e l s f o r t h e e f f e c t c f s i z e and d e n s i t y on g r c i t h . The * , * * , * * • i n d i c a t e a s i g n i f i c a n t l i n e a r r e l a t i o n a t t h e . 0 5 , .01 and .001 l e v e l s r e s p e c t i v e l y . ,  EXP  NET1 ME UN S I Z E BENS  NETW MEAN S I Z E DENS  "N ET D MEAN SIZE DENS  I  . 104 . 112 .12 1  . 145 .003  .040 .007  II  .174  .4 07  .242  .849 .106  .076  . 122 .008  III  .065 . 533 .578  .168  .084 .541  .056  .532  IV  .040 .492  .022  .536 .817  .018 ,457 ,96 2  NETWM MEAN S I Z E DENS  NETWS MEAN SIZE DISS  1.28 .041 .00 0  0.48 .070  0.81 .156  II  1.79 .023  .003  0.65 ,007 .001  1. 13 . 103-.€06  III  1.24 .036  .315  0.21 -450 .032  1. 03= .028 .5 70  IV  2.06 .014  .002  0.72 .000- .098  1. 35 .259 .000  EXP  PERL MEAN S I Z E DENS  EERW MEAN S I Z E DENS  I EE D MEAN S I Z E DENS  I  - 012 . 00 3-,252  .047 .009- .443  .027  .002  II  .032 .goo-- .65 3  .07 5 .847 .117  .047  .025 .023  III  .011 . 196 .318  i 0 4 8 .023 .2 58  .033 .519  IV  .007  .006  .012  EXP  PEEWT MEAN S I Z E DENS .  PERWM ME AK SIZE DENS  FIEWS MEAN S I Z E DINS  .111 .000  .003  .138  .021 .294  .102 .000  II  .135 . 0 00 .083  .154  .008  III  • 081 .003  .044  .249 .103  EXP I  I  IV  *  .012  **  *  .623  NETWT . ME UN SIZ E DENS  ** *  *** **  *** ***  T .597  *** *** if  .564  **  *  .277  ***  .271  .005  *** * *  .545 .7 66  **  **  .117  ,003  ** **  *  .911  .000  *** **  *  ***  .126  **  *  .010  **  **  ,S56  .436 .97 2  ***  .006  **  . 000 .140  ***  .099 .003- .606 •4*  .125 . 005 =.002  ***  , 139 .674  .720  .120 .005  ***  .000  ***  144  Table  27.  Non-linear l e a s t squares f i t (LSF), using a g r i d s e a r c h method ( B e v i n g t o n , 1 9 6 9 ) , o f the p r o d u c t i v i t y , P (ug C/ug C H L A / h r ) , v e r s e s l i g h t , E AE ( l y / m i n ) , d a t a f o r b o t h t h e •TA NE * and •SHITE' h y p e r b c l i c m o d e l s . The p a r a m e t e r A (1) c o r r e s p o n d s t o PMAX and r e p r e s e n t s t h e n a x i a um net p r o d u c t i v i t y (S «as s e t t c z e r o f o r the c a l c u l a t i o n s ) . The s u b r o u t i n e p a r a m e t e r s a r e d e s c r i b e d a t t h e end o f t h e T a b l e . 8  e  I . TANK  MODEL P = PMAX*TANH(S*PAS/PMAX) B  - B  fe  1. P ^S I - 14 DEG,(BLOCK),NAIEHA=2,N=7,P.NET ALPHA,S = 1 1 . 8 3 X(I)= 0.043 0. 103 0. 154 0.231 0.317 0.377 0.488 Y(I) = 0. 320 1.030 1.650 2.500 3.220 3.250 3.380 YFIT(I) = 0. 506 1. 182 1.705 2.372 2.S32 3.217 3.569 NPTS= 7 NTEBMS= 1 MCDE= 0 A(1) INITIAL= 3.900 FINAL= 3.986 DELTAA(1) I N I T I A L = 0.050 FINAL= 0.033 SIGMAA(1)= 1.219 CHISQB= 0.039 AVG Y ( I ) = 2.193 GAMM A= 0. 1966 1 GAMMAM= 0.08S66 2. P VS I - 16 DEG,(BLCGB),NALFHA=2,N=7,p.NET ALPHA,S = 21.83 X(I)= 0.043 0. 103 0. 154 0.231 0.317 0.377 0.488 Y(I)= 0.200 1.510 2. 150 3.960 6.050 5. 190 5.830 YITT(I) = 0. 932 2. 160 3.083 4.205 5.076 5.487 5.952 NE1S = 7 NTEEMS= 1 MODE= 0 A(1) INITIAL= 6. 200 FINAL= 6.392 DELTAA (1) I N I T I A I = 0.050 FIHAL= 0.067 SIGMA A (1) = 1.012 CHISQB= 0.588 AVG Y (I) = 3.556 G AMMA= . 2. 93949 GAMMAM= 0.82670 3. P VS I - 18 DEG, (BLOCK) , N AI PH A=2 , K= 7, P.NET fiIFHA,S = 25,83 X(I)= 0. 043 0. 103 0, 154 0.231 0.317 G.377 0.486 Y(I)= 0. 180 1.730 2.680 4.590 6.650 7.260 7.5 10 YFIT(I)= 1, 104 2.578 3.714 5. 156 €.356 6.961 7.70 0 NPTS= 7 NTEEMS= 1 MCDE= 0 A(1) INITIAL= 8.500 FINAL= 6.553 DELTAA (1) I N I T I A L = 0.050 FINAL= 0.0 17 ' SIGMA A (1)= 1.200 CHISQB= 0.667 AVG 5 (I)= 4.400 G A M M A= 3.33268 € AMM AM= 0.75743  145 4. P VS .1-20 DEG, (BLCOH) ,NALPHA*2,N=7,P. SET ALEHA,S = 3 1 . 8 9 X(I)= 0. 043 0. 1.0.3 0. 154 0.231 .0.3 17 0. 377 0.48€ Y(I)* 0.150 -1.340 3.6S0 6.010 7.350 6.780 8.66G YFIT(I) = 1.362 3. 165 4.53 1 6.217 7.557 8.203 8.952 NETS* 7 NTEBM S = 1 MODE* 0 A(1) I N I T I A L * 9.200 FINAL* 9.707 .DE-XT A A (1) I N I T I A L * 0.050 FINAL* 0.167 SIGMAA ( 1 ) - 1.071 CHISQB= 1.187 A VG Y (I) = 5.169 GAMMA5.93326 G A M M A R= 1. 14795 5. P VS I ~ 20 PEG,(POST-BLOOM),NALPBA=2,N=7,P.NET ALPHA,S •= 31.17 X (I)= 0. 043 0. 103 0. 15 4 0.231 0.317 C. 377 0.466 Y(I)= 0. 160 2.030 3. 230 5.950 6.760 7.090 9.000 YFIT(I) = 1. 330 3.074 ,4.371 5.92 1 7.096 7.635 6.228 NPTS= 7 NTEBMS*' 1 MODE* G A(1) I N I T I A L * 8.700 FINAL* 8.753 DELTA A (1) I N I T I A L * 0.050 FINAL= 0.017 SIGMftft(1) = 0.982 CHISQE* 0. 953 A VG Y <!)= 4.889 GAMM A* 4.76638 GAMMAM* 0. S75 01 6. ,P VS I - 18 DEG, (POST-BLOOM) ,NALFEI=2,N=7,E.NET ALPHA,S = 24.50 X(I) = 0. 043 0.103 0.154 0.231 0.317 0.377 0.486 Y (I)* 0. 130 1.600 2.690 4.610 5.760 6.640 6.640 YFIT(I)= 1.047 2.432 3.482 4.779 5.811 6. 310 6.886 NPTS= 7 NTEBMS* 1 MODE* 0 A(1) I N I T I A L * 7. 200 FINAL* 7.474 DELTAA (1) I N I T I A L * 0.050 FINAL* 0.083 SIGMAft(1) = 1.068 .CHISQB* 0.460 A VG 1 ( 1 ) - . 4.039 GAMMA* 2.30191 GAMMAM* 0.56SS8 7. P VS I - 16 DEG, (POST-BLOOM) ,N ALFBA=2, N=7, P. N E3 ALFB A ,S = 20.50 X(I) = 0. 043 0. 103 0. 154 0.231 0.317 0.377 0.486 Y (I) = 0. 120 1.350 2.620 3. 780 5.020 4.S1G 6.010 YFIT(I) = 0.876 2.034 2.912 3.994 4.853 5. 268 5.747 NETS* 7 NTEBMS* 1 MCDE= 0 A(1) I N I T I A L * 6.200 FINAL* 6.229 D EXTAft(1) I N I T I A L * 0.050 F I N A L * 0.017 SIGHAA(1)* 1.080 CHISQB* 0. 26 4 AVG Y ( I ) = 3.430 GAMMA* 1. 31836 GAMM AH* 0. 38436  146  I I . SMITH MODEL: P = PMAX*S*PAB/(SQET ( (EMAX**2) • ( ( S * P A B ) * * 2 ) ) ) - K 8  1. P VS I - 14 DIG, (BLOOM) , NALPHA=2,N=7,E. NET ALP HA, S = 11.83 X(I) = 0.043 0. 103 0. 154 0. 231 0.31 7 0.377 0.488 Y(I) = 0. 320 1. 030 1.650 2.500 3.220 3.250 3.380 YFIT(I) = 0.506 1.178 1.695 2.351 2.9 10 3. 207 3.605 NETS= 7 NTEBM S= 1 MCDE= 0 A(1) INITIAL- 3. 900 FINAL= 4.615 DELTAA(1) I N I T I A L - 0.050 FIN AL= 0.233 SIGMAA(1) = 1.501 CHISQB= 0. 046 AVG Y LI) = 2.1 S3 GAMMA= 0.22868 GAMMAM= 0.10429 55  2.,P VS I - 16 DEG,(BLOOM),NALPHA=2,N=7,P.NET ALPHA,S = 21.83 X(I)= 0. 043 0. 103 0. 154 0.231 0.317 0. 377 0.488 1(1) = 0. 200 1.510 2. 150 3.960 6.050 5.190 5.830 YFIT(I) = 0. 931 2.149 3.053 4. 148 5.020 5.458 6.017 NETS= 7 NTEBMS= 1 MCEE= 0 A(1) INITIAL= 6.200 FINAL= 7.292 DELTAA(1) I N I T I A L = 0.050 FIN AL= 0.367 SIGMA A (1)= 1.253 €HISQE= 0. 592 AVGY(I)= 3.556 GAMMA= 2. 96142 GAMMA K= 0.83286 3. P VS I - 18 DEG,(BLOOM),NALPHA=2,$=7,E.NEl ALPHA,S = 25.83 X(.I)= 0. 043 0. 103 0. 154 0.231 0.317 0.377 0.486 Y(I)= 0. 180 1.730 2.680 4.590 6. 650 7.260 7.510 YFIT(I)= 1. 104 2.568 3.688 5. 103 6. 295 6.923 7.75S N£TS= 7 NTEBMS= 1 MCCE= 0 A(1) INITIAL= 8.500 FINAL= 9.844 DELTAA (1) INITIAL= 0.050 FINAL= 0.450 SIGMAA (1)= 1.496 CB.ISQB= 0. 664 AVG Y ( I ) = 4.400 GAMMA= 3.31827 GAMMA«= 0.75415  B  147 4. ? VS I - 20 DEG, (SLOCK) ,NALEHA=2,N=7,P.NET ALPHA, S = 3 1 , 8 9 X(I) = 0. 043 0. 103 0, 15 4 0. 231 0,317 0.377 0,486 Y(I) = 0. 150 1.340 3.690 6-010 7.350 6.780 8.860 YFIT(I) = 1.361 3*150 4.49 1 6. 137 7.472 6.153 S.033 NPTS* 7 NTEBMS* 1 MODE* 0 Ml) I N I T I A L * 9.2G0 FINAL* 11.093 DELIAA (1) I N I T I A L * 0.050 FINAL* 0.633 SIGMAS (1)= 1. 330 CHISQB* 1.167 A VG Y ( I ) = 5.169 GAMMA* 5. 83592 GAMMAM* 1.12912 5. P VS I - 20 DEG, (EOST-BLOGM) ,NAIFHA=2,N=7,F.NET ALPH B,S = 31. 17 X(I) = 0. 043 0.103 0. 154 0. 231 0.317.0.377 C.486 Y(I) = 0. 160 2.030 3. 230 5.950 6.760 7.GS0 9.000 YFIT(I) = 1.328 3.055 4,321 5.828 7.003 7. 583 8.313 NETS* 7 NTEBMS* 1 HCDE* 0 A(1) I N I T I A L * 8.700 FINAL* 9.927 DELTA A (1) I N I T I A L * 0.050 F I N A L * 0.417 SIGMA A (1) = 1.216 CHISQB* 0.879 A VG Y ( I ) = 4,685 GAMMA* 4.39499 GAMMAM* 0.89903 6. P VS I - 18 DEG,(POST-BLOOM),NALFHA=2,N=7 E,NET ALEHA S * 24.50 X(I) = ' 0.043 0. 103 0. 154 0. 23 1 0. 317 0,377 0.486 Y(I)= 0. 130 1.600 2.690 4.610 5.760 6. 640 6.640 YIIT(I)= 1. 046 2.420 3.452 4.719 5.747 6.273 6.95/ NPTS* 7 NTEEMS* 1 MODE* 0 A(1) I N I T I A L * 7. 200 FINAL* 8.546 DELTA A (1) I N I T I A L * 0.050 FINAL* 0.450 S1GMAA(1)= 1.335 CHISQB* 0.450 A VG Y ( I ) = 4.039 GAMMA* 2. 25042 GAMMAM* 0.55723 #  7. P VS I - 16 BEG , (EOST-BLOOM) ,NALEBA=2,N=7,P.NET ALIHfl,S= 20.50 X(I)= 0. 043 0. 103 0. 154 0. 231 0. 317 0. 377 C,488 Y(I)* 0, 120 1.350 2. 820 3. 760 5.C20 4.S1C 6.010 YFIT(I) = 0.875 2.024 2.866 3.944 4.801 5. 238 5.802 NETS* 7 NTEBMS* 1 MODE* 0 A(1) I N I T I A L * 6.200 FINAL* 7.123 DEI-T A A (1) I N I T I A L * 0.050 FINAL* 0.30C SIGMA A (1)= 1.319 CHISQB* 0.251 A VG Y (I) = 3.43 0 GAMMA* 1. 25428 GAMMAM* 0. 36568  ma  DESCRIPTION OF PABAHETEIS IN TEE NGN-LIK'E A E LSF SOEEOuIINE S - I n i t i a l s l c c e , ALPHA R  -. R e s p i r a t i o n  (Input Parameter)  (Input Parameter)  X - Array of data p t s . f o r indep var Y - Array of data p t s . f o r dep var  (PAB)  (P ) B  NTERMS - No. o f parameters MOLE - Determines method of wgting LSE A - Array of parameters o f be estimated (A=1; A (1)=PMAX) DELTAA — Array o f increments f o r parameter(s) A SIGMA A - Array o f S t . Dev. f o r para meter(s) A YFTT - Array of c a l c u l a t e d values c f Y CHISQB - Seduced C h i Square f o r f i t AVG - Average Y (I) value GAMMA — Sum of ( (YFTT (I)-Y (I) ) **2) GAMMAM - GAMMA/AVG  Figure  incoming seawater-£  1.  Experimental  facilities  f o rExperiment 1  fiite^U7-SEAWATER RESEVOIR fluorescent lamps 'oooooooo—00000000  -inflow tube  HOLDING TANKS FOR GRAZERS  Ioutflow tube  EXPERIMENTAL TANKS jn situ sampling tubes  1 32.  inflow pvc tubing  V I E W  VS  EXPERIMENTAL TANKS  outflow. S U R F A C E  W  ®  VALVES  F i g u r e 2.  D u p l i c a t e tank systems  TANK. A  f o r Experiments 2 and 3  TANK  B  F i g u r e 3.  S i d e view o f t h e e x p e r i m e n t a l f a c i l i t i e s f o r Experiments  2 and 3  SEAWATER RESEVOIR plywood c o v e r ,  IPS V PRIMARY PRODUCTION TANKS plexiglass cover.  S C A L E : 1cm= 0.3 metre SCALLOP TRAYS or OYSTER CULTCH •  VALVES  SIDE VIEW  Figure 4.  Side view of the experimental f a c i l i t i e s for Experiment 4.  SEAWATER RESEVOIR  plywood cover,  V ^X^e.  PRIMARY PRODUCTION TANKS plexiglass cover.  SCALE: 1cm= 0.3 metre SIDE VIEW VALVES $  H-  punp ®  fo  F i g u r e 5.  Two-stage c u l t u r e systems f o r Experiment  H2  H3  CONTROL  .  5  ACCLIMATION  153  F i g u r e 6.  N i t r a t e c o n c e n t r a t i o n d u r i n g Experiment  IA  F i g u r e 7.  N i t r a t e c o n c e n t r a t i o n d u r i n g Experiment  IB  • F i g u r e 8.  P h y t o p l a n k t o n s t o c k d u r i n g Experiment IA  _ . . _  SURfflCE MID BOTTOM  SAMPLING  If \ V >' f \\/ 0.0  5.0  It  N  I S 10.0  T  T 15.0  I  20.0  25.0  30.0  MY  s 35.0  TIME (Dfir)  40.0  I  45.0  —I  50.0  55.0  60.0  65.0  70.0  Ln ON  F i g u r e 9.  P h y t o p l a n k t o n s t o c k d u r i n g Experiment IB  sw m orsct BOTTOM  a  SWUNG 0R1  A / \ / \ / \ / \ / \  //V 0.0  T S  s.o  10.0  IS.O  20.0  ~ \ /  2S.0  30.0  V /  35.0 TIME (DAT)  40.0  —1— 45.0  n — SO.O  -1— S5.0  60.0  6S.0  70.0  Figure 10.  Primary productivity during Experiment IA  PRIMARY PRODUCTIVITY - IA  S U R F A C E W D  eons* +  B  D E T R I T U S S R H P L J N G o a r  oo  on  30.0 I  [00.0 I  PRIMRRY PRODUCTIVITY (uG 150.0 203.0 iSO.O I  I  l_  1 4  C/LTTRE/HR) 330.0 1  350.0 I  .  n  410.0 1  ^ n n <50.0 1  500.0 1  OQ  c  H ro  •  v. «^ T3 l-f O Cu  c o  rt H" <!  I  c  v!  •V  3 CN  m ro S  ro 3 ts  r  y  171  s-  6ST  F i g u r e 12.  Primary  productivity  ( s t a n d a r d i z e d ) d u r i n g Experiment IB  F i g u r e 13.  S o l a r r a d i a t i o n d u r i n g Experiment 2  162  Figure  q  °i^>  £^  14.  T e m p e r a t u r e d u r i n g E x p e r i m e n t 2A  TTo  iTo  jjTo  2E0  TIME  (DRY)  iTo  SE©  -  «tio  _____  4Z0  ~sl.o  163  Figure 15.  Temperature during Experiment 2B  JNFLOV SURFACE  HID Know OUTFLOW  " °  4  1  0  »'••  *.o  a.'.o  ^To TIME (DAY)  "ik  sU  4.'o  ^  164  Figure 16.  N i t r a t e c o n c e n t r a t i o n d u r i n g Experiment  TIME (DRY!  2A  165  166  Figure 18.  Phytoplankton stock during Experiment 2A  5URFRCE HID B3T10H OUTFLOW x  SRMPL1NG DRY  A  /\  •V.  ho  * * * * x  to  ii I »  11.0  "*~T—5  r~5 Z/.o  i ) (DAY) TIME -  x  %.o  I  3/.o  34.6  — i  1  Figure 19.  P h y t o p l a n k t o n s t o c k d u r i n g Experiment  2B  168  Figure 20.  Oxygen concentration during Experiment 2A  a  T o  itlo  7^6  7£o  2/.*  i£o TIME  (DAY)  sLe>  <A.o  t/i-a  &).a  169  Figure 22.  Primary productivity during Experiment 2A  171  F i g u r e 23.  Primary p r o d u c t i v i t y  d u r i n g Experiment 2B  SURFACE  HID. B8TT0M  orioLU  rr  go  a.  x  SRMPUNG OflV  172  F i g u r e 24.  Si  Primary p r o d u c t i v i t y  (standardized)  d u r i n g Experiment  2A  173  Figure 25.  Primary productivity  (standardized) during Experiment 2B  SURFRCE MID BOTTOM  ll  cn".  X  SAMPLING DAY  CJ (JD  ll  —ri. C J  /'  ! i I i  » —.  co  i  i  .i' • /  C J  CY.  \ / U-\  :  _=  _  •—ICO  O.  I' *  ' /.'  i>  V "  1  v  I.O  *-0  //. O  /*.©  i/.O  34>4>  31.- o  ~' 3ko  TIME (DAY)  tf/.o  ti&.e>  174  175  F i g u r e 27.  Temperature d u r i n g Experiment 3A  imov SURFACE HID BOTTOM OUTFLOW  A  °  I  ^  0  77Z  " °  ~J^o —  —  zi^  TIME" (DAY)  176  Figure 28. Nitrate concentration during Experiment 3A  S-4  4  A  UJ  cc  32' UJ  •—  cc cc  IO  x x x x x x x x x x x 4.0 11.©  iL.o  jiro  it* TIME IDRY)  ai'.o  _  imov SURFRCE HID BOTTOM OUTFLOW  n  SAMPLING DRY (5.H.B)  177  F i g u r e 29.  to  P h y t o p l a n k t o n s t o c k d u r i n g Experiment  3A  Figure 30.  Oxygen c o n c e n t r a t i o n d u r i n g Experiment  3A  179  F i g u r e 31.  Primary p r o d u c t i v i t y  d u r i n g Experiment  3A  HID BOTTOM  TIME (DRY)  180  F i g u r e 32.  -  •  '  '  •  «  4-»  Primary p r o d u c t i v i t y  x  X  i x  n.o  x  I x  /Co  XT  xto  (standardized)  T  X*-0  |  JI.O  d u r i n g Experiment  1  ii.0 TIME (DflYJ  1  IHO  3A  i  .  *'o  181  Figure 33.  Solar r a d i a t i o n during Experiment 3B  182  183  Figure 35.  N i t r a t e c o n c e n t r a t i o n d u r i n g Experiment  3B  184  F i g u r e 36.  P h y t o p l a n k t o n s t o c k d u r i n g Experiment  3B  SURFRCE HID BOTTOM CUTFLOW SAMPLING DAY (5.M.B)  *7*  fc.o  (1.0  l~4>  i TIME  (DAY)  9>.o  SI*  186  F i g u r e 38.  Primary p r o d u c t i v i t y d u r i n g Experiment  3B  ml  SURFACE HID BOTTOM  SAMPLING DM  x  I  l.o  »  M  * >  £.o  *  ii  I  ii  ti-o  ii  I ic  lU.o  £  I ii  37-0  ii  T"  ii  2-L o  TIME (DAY]  J  ii  ?/.o  I  3C.0  I  I  *H-o  I  *U,.o  Si.o  Figure 39.  Primary productivity (standardized) during Experiment 3B  188  FIGURE  4-0  FORCING CONDITIONS DURING EXPERIMENT 4  SOLRR RROIRTION TANK  fl  im  B  FIGURE 4V  TEMPERATURE DURING EXPERIMENT 4  IHfLOV  °.  SURFRCt  BOTTOM  MID  OUTFLOW  TRNK R  190  FIGURE  NITRATE  M  DURING EXPERIMENT 4 IHFLoY SURFRCE  BOTTOM  MID  OUTFLOW  TRNKR  8,0  16.0  40.0  43.0  40.0  49.0  TRNK 0  TI^ARYS)  3  2  ,  0  FIGURE .43  PHYTOPLRNKTON DURING EXPERIMENT 4  192  CHLB'.CHLR RATIO  FIGURE W  DURING EXPERIMENT 4  SURFRC£  BOTTOM  MID  OUTFLOW  TRNK R  r~—i 16.0  '1ME? 1°DRYS)  1 32.0  1  1 j 40.0  r 48.0  4  TRNK B  A. T T T 24.0 32.0 TINE (DAYS)  40.0  43.0  FIGURE 4-5  CRROTENOID:CHLR DURING EXPERIMENT 4  SURFRCE HID  Bonem  OUTFLOV  TRNK R  vn 2P cc  ZZ CM Q N  o  z UJ ajg  C E •_ C C -  o o"  o.o  —r— 8.0  T IS.O  r  T  24.0  TIMbf  T  r  32.0  43.0  43.0  32.0  43.0  43.0  IDRYS)  TRNK B  a in"  I—tn" CC CC  51 X Q Z. UJ OH  cru o  • a 0.0 m  -] r 8.0  16.0  TIME?VDRYS)  FIGURE H  OXYGEN  DURING EXPERIMENT 4  INFLOW  SURFRCE  wnoH  HID  OUTFLOW  TRNK R  x  .Vvr  V"v f\/vsi/ :  0.0  -)—  8.0  T U  -°  TIH^ftflTS)  ,v;  40.0  48.0  TRNK B  0.0  e.o  43.0  — I —  49.0  195  FIGURE  GROSS PRIMRRY  47  PRODUCTIVITY  DURING EXPERIMENT 4 SURFACE  Q  ©  MID  anion _ o  TANK A A  1°. GJ  9  A  A  •  © ©g g _ _ ©  0  © m  A  m  O  ©  8  cn a  A  A<  1  — I  0.0  1  J——I  8.0  .0  16  T  ]  1 |  1  ^  R  1 Y  S  )  1  1  1  1  3*.°  1—  «.0  TRNK B _ ©  CC  1 °  d  CD  -  O  a "  B  g  ©  ®  •  A  B  A  CO  © CJo  ©~  A  _JCM  ©  a. " a  tn  _  o a  1  0.0  1  6.0  1  1  1  1  r——I  16.0 --jj^.O TIME IDRYS) 32.0  1  1  40.0  1  1  48.0  FIGURE 4 8  RESPIRRTIQN RRTE DURING EXPERIMENT 4  SURFACE  •  MID  Q  BOTTOM  A  TRNK R  8-  HffJ *—I  I—Q  CD  85  13  — i 0.0  1 e.O  1  A  A  _  m  CD  is *  6  1 16.0  o1 1 ^,.JA.Olr,  ,  TIME IDRYS)  1  1. m " 32.0  i  1 40.0  1  1 40.0  1  1— 48.0  TRNK B Io  l—a QCCM.  A  CL  _  UJ CC  A  .  e>  8 8  @  °  CD  A  °  g  H  0.0  T 6.0  1  1 16.0  1 ,  Tlir  1 Z4.Ci  1 „  TIME (DRYS)  A  1 32.0  1——r  48.0  197  FIGURE  W  NET PRIMARY PRODUCTIVITY DURING EXPERIMENT 4  SURFACE Q MIC  Q  BOTTOM  A  TRNK R  CDcV  ^ . aI *a  C3  T  "  1  *-°  1 i S  "°  •f  1  1  TJME  24  1  (DRYS)  1  r——i  3 2 - 0  m  1——r  '  Q  4  8  0  TRNK B  UJ  A  B "  •J  S S  T^  I 1  8.o  1  —1  ,e.o  ^ | om  1  B ! H 1 n H  I  ^  1  f t f i Y S )  1  1  1  „.„  1  1 m j t  1  198  FIGURE 5 0  NET"PRIMARY  PRODUCTIVITY  (DRILY) DURING EXPERIMENT 4 SURFACE  A  HID  ©  BOTTOM  _  TRNK R  o  LL)  CJD ' 01  A  LU  CD  Q  A  ffl  —r  a  1  |  8  A  1 .6.0  O  t CD 1  T  3  H  E  I  1 ^ 0  ©  D  f  l  S A ^ A r—i 32.0  Y  S  H  1  )  1 41.0  1  1— 43.0  1 1 40.0  1— 48.0  TRNK B V"  >CE  UJ CC  Cu CL -4 UJ —  a S jm fi_ aa 5* 0.0  —| fi.O  1  1 1 1 16.0 _.._24.0  »  u  1 _  •JHfcTftflYS)  §  m  1 n 32.0  199  FIGURE 5 \  GROSS PRIMRRY  PRODUCTIVITY  (DRILY) DURING EXPERIMENT 4 SURFACE Q HJO Q BOTTOM A  TRNK R A  01  cc  A  S A  B  A  © o GJ  CL.  a-  A  S  •  s  cn-'  CD  A  (D (D .  —I  r  1  1  i6.o  n  1  T ] M  ^  GJ  8  CD  *  A  r——r  1  1  <o.o  *.o  A f ) Y S )  1  1  «.o  TRNK B C3  &  a  CD  ft CJ  UJtr)'  *  ^CD *A  Q  A  A  ^ CD B  •  CD U  °  O  T  0  G)  GJ  A  CL.  cn  a  cc .  CD  o o"  1——I  1  1  ' -° 6  1  1  1  1  TJME^RYS) * -  1 3  1  *•*  1  1  1  200  FIGURE 5 2  GROSS PRIMRRY  PRODUCTIVITY  (STRNDRRDIZED) DURING EXPERIMENT 4 SURFACE GJ HID Q B0T1OH  A  «  TRNK R  CM  CC • X  x-  o  CD . Z3  ©  a  _  A  •  \  © o  ZD e  A  G3  _  _  ci-q  A  cc H CJD  o o  -i  1  0.0  1  8.0  1  1  i  1  1S.0 ^ ^ . 0 ^ ^  r  1  32.0  1  1  43.0  1—  43.0  TRNK B cc • _J<D. CJ C3 . ZD A  °  B  _  O  ©  _  A  & © CD  A  A  CO CD  §  o a  m  © a  m  a  H  CD  1  1  1  1 1B  1  1  1  '° 7i«?Ams>  1  1  1  1  1 4  U  201  FIGURE S 3  NET PRIMARY PRODUCTIVITY (STRNDRRDIZED) DURING EXPERIMENT 4  SURFACE HID QJ Q  o  TRNK R  CM  CC  x  Q  x CJ CD  ClgfJ 3  A  OL  I—  UJ  ^  9 6 | r  — i  1  M-  I  B 1  1  IM  T  I  5  1  I  |  « f t  c  1 B  Y  1  S  1  *.«  )  1  1  «•••  1—  «•«  TRNK B  EC CEflO. X  o  CD CD" Z3  Cu  o  »— UJ ZZ •  Ala 8 — i  1  1  8  $  1 UJ.  B & 1 1  J  f  E  1 « f t  1—n R  Y  S  )  1  1  1  1—  202  FIGURE-54 SURFACE  RESPIRRTIQN RATE (STRNDRRDIZEDJ DURING EXPERIMENT 4 a  o  HID B0T1OH  A  9  TRNK R  CM  cc _  CJ  3  m  m  & -J  _  8  A  UJ  r  B 1  i8.o 1  T1 ]ME  A  8  *Y1  A  A A  CD  A  GJ  »•1«»  DflY1 S)  1  -.o 1  1  r«  TRNK B  cc X V. CEO  x-;  M  CJ CD  =1  •v.  1  Oo C D ™ _  _  CE  op CL  cn . UJ cc  A  A "I  1  1  1  1  A 1  8 r— A —i  1  1——r  203  FIGURE 55  ESTIMATES OF ALPHAC DURING EXPERIMENT A  SURFACE MID © BOTIOM A E  TRNK R  cn x v, •  CJ  V. -  CJ  28-  u cc . a. 0.0  A  a.o  T  '  l 0  8 -A  T  n TIH^ftflYSJ  ^  40.0  43.0  43.0  43.0  TRNK B  CC X  5CD V.  .  CJ CD . 0  3sCJ  ex . X D_  cro. o 0.0  i —  8.0  ft, A 16.3  @ i  TIME ftRYS).  •  32.0  204  FIGURE 5 4  ESTIMATES OF ALPHAG DURING EXPERIMENT A  SURFACE  Q  MID  o  TRNK R  J8v." cc IT v. •  ®  GI  a A  g  8  X  E  o.»  —I  A  O  n>  •  Qu  cro  A  r . . .  1  ©  g  m o  GJ  g  1 1 .6.0 m  i  ^  1  1 m  S  TRNK B  1 * . »  i  1  1  1  — i — «  £ 7E0.0  '—  cr x \  cc  •  ZJ°.  o§CD Z3 v. .  u  A A  ©  8 A  CD <X X D_  CD ©  J  H  fife 0.0  a  ©  CD°.  ©  ©  ©  m  GJ Q GJ —i— 6.0  A  ' -° T I M E R S ) -° 6  33  40.0  I— 43.0  Figure 57.  Coulter counts on Day 6 of Experiment 4A  Figure  59.  C o u l t e r counts on Day  15 o f Experiment  4A  COULTER COUNTS 417 15 SURFRCE— MID _ _ _ BOTTOM- _  Figure 61.  Coulter counts on Day 27 of Experiment 4 A  COULTER COUNTS 417 27  FIGURE 6 1  FORCING CONDITIONS DURING EXPERIMENT 5  211  FIGURE  TEMPERATURE  43  DURING EXPERIMENT 5  INFLOW  SURFRCE  B8T10H OUTFLOW  KID  * 8n  oT  TRNK R  r  r  1  n 16.0  1  r-—i  1 r-—i 32.0 43.0  1  1 1 43.0  212  FIGURE  64IHFLW SURFACE HID  NITRATE  DURING EXPERIMENT 5 BOUGH OUTFLOW  213  FIGURE 65  PHYTOPLANKTON STOCK DURING EXPERIMENT 5  SURFACE MID  °. Si  cf 1  BOUGH .....  ......  TRNK R  CUTflCfl  214  FIGURE  CHLB:CHLR RRT10 DURING EXPERIMENT 5  U  SURFACE HID  WHOM OUTFLOV  TRNK R  18.0  .  1 l~ 24.0  TIME IDRYS)  32.0  —I  40.0  43.0  TRNK B  A 16.0  r „. 24.0  TIME IDRYS)  • tjXl.  32.0  40.0  "1— 43.0  FIGURE LT  CfiROTENOID:CHLfi RRTIO DURING EXPERIMENT 5 Barron  HID  OUTFLOtt  TRNK R  1  -I  8.0  0.0  43.0  TRNK B  i  i  1  1  .0  16  1 T l f E  1  «. ° (  1 D f i r g )  1  SM  1  1  1  «.  0  1  «.0  1  FIGURE 4 8  OXYGEN DURING EXPERIMENT 5  FIGURE Ll  GROSS PRIMARY  PRODUCTIVITY  DURING EXPERIMENT 5 SURFACE [TJ MID © amon A  O.JO o.25  TRNK  8_  FLUSHING RATES 0.50 . i.oo .  R  CD  S  ID CD  a-  D  g  o § CD  ej  o"T  I  I  1  1 L 6  -°  1  1  1  1  UIC'VDRYS;  » O  a  IS '  1  T  t  A A  m  1  * "  7RNK B GJ  r-  E  st  ©  *•  o GJ  A o i  8  T  Gl i  r—  1  ,6.0  1 U(1  1  £Y  1 DRVS)  T  »•»  f-  1  «•».  1  nr  218  FIGURE 70  GROSS PRIMRRY  PRODUCTIVITY  (DAILY) DURING EXPERIMENT 5 SURFACE a  MID  FLUSHING  o  BOTTOM  0.10  0.50 ,  0.23  A  RATES  1.00  .  TRNK R  ex CD  B 8  H  A  CD  A  Q A CJ  CD  ^  ca  B I  "I  1  i  1  1—;  i  1  1  ,S.O  1  i  —  i  mM  «.»  TRNK B  93  >a  § ?  6  cc  S  2  cn~ cn  *  »  S  A  CD A Q  ro i  ~r~ a.o  i .  i 16J  i T  J  i H  E  «  f  i t  f  i  Y  S  1 )  ».•  1  1  1 «  1—  1  FIGURE 71  RESPIRRTION RRTE DURING EXPERIMENT 5  SURFACE  FLUSHING  N  RATES  HID  ©  0.10  0.50 .  WTTOH  A  0.25  1.00 .  2  _TRNK_ R_  W o i  1  -  -  i  1  3.0  1  5  m  fl  g  °  1  16.0  *  .  1  \  0  m  1  O  •  1  ©  r  1  «•* ^ «  ^Yn,^ *•» TflNKB  • * x  ®  ®m  a  A  •21 H . ".  8  -°  16  ' ft ~  fi  A  A  * T I I -° TIHtTftflYS)  S. o  B  *  D  «  8 i  ~ .  _  A  u  tp  —  , 4 ,  —  -°  i  * ,  i  *-  — 0  ,  —  220  FIGURE 72  NET PRIMARY PRODUCTIVITY DURING EXPERIMENT  SURFACE GJ KID  FLUSHING RATES ffl  BOTTOM  5  A  0.10  0.50 „  0.Z5 _  1.00  TRNK  R  ©  cc° CD ©  >M3  CScV  8  i— UJ  a o  ©  oo.  T  1  i<?  1  1  1  m  «•»  1  1  ms}  TANK  1  1  «-»r 1  B  UJ cc ">vO  ©  CD CM J  2  8 A  Q"M. .S I  A ft  ©  D  ©  I dT 0.0  u  ®^  1  6.0  A  Q  A  e  m  aa  a1  1  16.0  1  1  „  AA  1  *  ^ g  e ft g  FFL  1  24.0 32.0 IDflYS)  TIME  a  1  A  1  40.0  A  1  1  46.0  «•»  221  FIGURE 73  NET PRIMARY  PRODUCTIVITY  (DRILY) DURING EXPERIMENT 5  SURFRCE a  FLUSHING FATES  MID  Q  O.JO  O.SO  BOTICH  A  0.23  1.00  TRNK R O  tu cc A  0J  o  8  UJ  6  a 1  1  1  8.0  1 GJ 1  16.0  7JMF  A  1  fY  1  1  DRYS]  32.0  1  1  1  1  40.0  1  41.0  TRNK B  UJ CC  GJ  A  8 .  0  ,GJ  I  UJ  A  ffl  Q  2.  E  •  ffi  s  .. H  m  A  ?  -  *  «  s  A  a  . ffl  @  E  GJ  o  1  0.0  e.o  1  1  J——I  1  1 6 . 0 ^ftflYS)  1 3 2 , 0  1  1  1  1 43  *°  1  222  FIGURE  74  GROSS PRIMARY PRODUCTIVITY (STANDARDIZED) DURING EXPERIMENT 5  SURFACE QJ MID © BOTTOM A  RUSHINS RATES 0.10 0.50 . 0.23 1.00 .  TANK R •  CC  ©  • X  ©  X "  o  to  ZO  A  CD . Z3  ©  fi  © o  ©  © ©  Gl  B  ©  m  A  ©  til ©  CD  0 1  E  A A  CO A  —P—  0.0  T  8.0  T  T  40.0  ^IHfcfftflYSl ^ °  43.0  TRNK B cc • x  A  ©  CJ  8  .  CO v.  m  i  s  3  ©  A  CO CO Q  © ©  GJ  1  ©  © A  CD fi  O  8  CO  5 0.0  i —  6.0  1  HI ,6  -°  1——!~  TiNTftms)  1  1  1  1  1  *  1 J  223  FIGURE 7 5  RESPIRATION RATE tSTRNDRRDIZED) DURING EXPERIMENT 5 FLUSHING RATES  SURFACE GJ MID Bonctf  0.50 . 1.00 .  0.10 0.25  CO A  TRNK R  Q TT. CM  CC J  aa  03  u  A GI  a --l Q  cc* M  UJ  ^  0  ffl @  cc  &  CD ®  0.0  I— 8.0  o i 1S.0  n i  H 1  £3  CD  L ._  24,0  1  TIME iDRYSJ TRNK B  @  43.0  43.0  r  1  32.0  cc J  eta xHCJ  CD Z3  •  CJ)CM. 3 " *  CX ..  6cj"  0.0  E GJ  O  A  m  ~r e.o  a g o 9 a?  Q 16.0  T  T  43.0  T  43.0  224  FIGURE  %  'NET'PRIMARY  PRODUCTIVITY  (STANDARDIZED) DURING EXPERIMENT 5  SURFACE HID  n ©  BOTTOM  O.JO  FLUSHING RATES •'; 0.50 ,  0.25  A  l.OD  .  TANK A  a v.  CM  CC IT  03  - J - *  a  X  CJ CD  9°  a  <JeJ. CD  ZD  ©  ©  a, m  'a  Cu  o  A ©  a  ©  UJ  ~ ~ i —  8.0  A  0  A  a 0.0  m  40.0  1S.0  48.0  TANK B  5** XX  txajJ o CD  "  CJw.  2  A*  ©  A  CUtD"  © 3  UJ  ©  0.0  ©  e.o  16.0  1  © A  S A  g  „  24.0  1  A  ©  (fi 1  TIME (DAYS)  1 —  32.0  40.0  49.0  225  FIGURE 77  ESTIMATES  GF RLPHAC  DURING EXPERIMENT 5 SURFACE MID  FLUSHING RATES 0.10 . 0.50 .  E  ffl  0.23  TRNK  9  1.00 ,  R  X  5§J CO  a  1  3sH GjCD  CJ  cr.  a  1  ,0  r  1  8.0  -i  16.0  1  1—"i  TRNK  r  43.0  48.0  B  X  55C9 CJ  co°.  38" o  £• cu 0.0 a k  'A  03  i 6.0  16.0  S 24.0  TIME  N  IDRYS)  A g  32.0  ffl A CD ffi  i g i 40.0  r  48.0  226  FIGURE 78  ESTIMATES OF ALPHAG DURING EXPERIMENT 5  SURFACE rrj MID ©  FLUSHING RATES 0.10 0.50  BOTTOM A  0.23  1.00 !  ;  TRNK R  9_  CC  5gJ  CQ  CJ  &1 A ^A  _ Ag  *  CD  o  A  0?  A  &  A  CD  © 0  m  .  ©  A  .  5  1  1 43.0  CU  1 S.O  — I  1  1 18.0  1 T  I  M  ^  1 0  L  "1 3  R  Y  S  1 3 .0  J  2  1  1— 43.0  TRNK B  Jh dg-l cc X  5°. C5  A  \ CJ  cs<=> , ES-i  S  A A A A  M  — I  1  6.0  1  1  !S.O  A  0 0  CD  ah  A O  A  1 T  1  ^  f  t  f  0  0 0 1 i  Y  S  A 0 0 1  )  33.0  A 0  A  0  0  1  0 0  A ©  1  41.0  1  1  «.0  VOLUME PER ML (XlO" ) 0.0  100.0  200.0  300.0  430.0  500.0  600.0  700.0  800.0  900.0  1000.0  Figure 80.  Coulter counts on Day 9 of Experiment 5A COULTER 517  COUNTS 9  SURFRCE  MID BOTTOM. _ _ _ _  —i—  —\—  —i  —I  PARTICLE SIZE (M)  ~ I —  18-o  Figure  81.  C o u l t e r counts on Day  21 o f Experiment  5A  COULTER 5 1 7 2 1  COUNTS  SURFRCE  MID  BOTTOM  . _ _  Figure 82.  Coulter counts on Day 24 of Experiment 5A  COULTER COUNTS 517 24 SURFRCE__ HID BOTTOM ___  Figure 83.  Coulter counts on Day 36 of Experiment 5A COULTER COUNTS 517 36 SURFRCE  MID BOTTOM-  :  .  Figure 84.  Coulter counts on Day 39 of Experiment 5A  COULTER COUNTS 517 39 SURFACE MID _ . _ ._ BOTTOM  a  a-  2= a  UJ  S i  /  —\— 34.  I—  I  S.7  -r PARTICLE SIZE ( A )  1—  \s.o  30.. b  tsj  co  Figure 85.  Coulter counts on Day 6 of Experiment 5B  COULTER COUNTS 527 6 SURFACE MID _ _ _ BOTTOM : _.  0  s-  UJ Q-  UJ  ».f  i  7-»  PARTICLE  ii.a SIZE(xc)  lY.o  N5  Figure 86.  Coulter counts on Day 9 of Experiment 5B  Figure 87.  Coulter counts on Day 21 of Experiment 5B  COULTER COUNTS 527 21 SURFACE MID _ . . ._ BOTTOM.  2=c c _ UJ  Figure 88.  Coulter counts on Day 24 of Experiment 5B  COULTER COUNTS 527 24 SURFRCE____ MID . . . . . . . BOTTOM  cc"  1  UJ Q.  UJ  i — i-fe  ~1— *-7  —r  -  i  N3  18-0  11.3-  PARTICLE SIZE  C*t)  2Z-L  38.  f  OJ  8  Figure 89.  Coulter counts on Day 36 of Experiment 5B  COULTER COUNTS 527 36 SURFRCE  MID _ BOTTOM  LU CL. LU  >  •  si  a-8  —r i-t,  1  *.S  —I  1—  9,1  7.1  1  q.o  :  r—  11.5  PARTICLE SIZE  1  ;*.3  1  (S.e  1——  r —  MJJT  NJ  Figure 90.  Coulter counts on Day 39 of Experiment 5B  COULTER COUNTS 527 39 SURFACE  MID BOTTOM  FIGURE "11  239  TEMPERATURE IN TVQ-STRGE OYSTER CULTURES  WIN  TANK ]  TRNK 3  TANK 2  TANK 4  EXPERIMENT  ca  1  CXPER1MEIT  8-  II  CJCVJ  i CO UJ  I  CO UJ  • -  a  UJ°-  •  UJ . Q  §2-  i— ac  cc UJ Q.  §3-  o o  —I  0.0  1  2.0  1  1  1  4.0  TIME tDRYS) EXPERIMENT  1  6.0  III  TIME tDRYS)  1  on  0.0  1  1  2.0  1  1  4.0  1——i  TIME (DRYS) EXPERIMENT  IV  6.0  1  FIGURE °iZ IHFLOV  CHLOROPHYLL R IN TVD-STRGE OYSTER CULTURES _  I  TAW 3  TRW 2  TRW 4  TRW  240  241  FIGURE  <B  OXYGEN LEVELS IN TVD-STRGE OYSTER CULTURES  INFUW TANK 1 TANK Z  q id.  TANK 3 TANK 4  EXPERIMENT  EXPERIMENT U-  I  a  a LU CC  UJ  CDo SOLD  co  -J.  CD  fe  a a'  0.0  a-.  2.0  i  1  TIME ('8flYS) 4  EXPERIMENT  o  1——r~ G  a"0.0  '°  2.0  1  1  1  4.0  r— 6.0  TIME (DAYS) EXPERIMENT IX  m.  a cvi.  a UJ cc  UJ cc  CDo  CDo  UJ CO >-  UJ  d'  ~~i  to >-  0.0  •  0.0  T  20  1 1 1——r 60 -— MrTIME (flfWS)  1  242 Figure 94. Growth of oysters as a function of DENS i n the two-stage culture ( DENS 1 (ggj DENS 2 DENS 3)  EXP 1  *7  EXP 2  EXP 3  EXP 4  EXP 1  EXP 2  EXP 3  EXP 4  EXP 3  EXP 4  EXP1  EXP 2  EXP 3  EXP 4  10  w  O DC  0 4.0  UJ  Z  5  V  EXP1  S3  EXP 2  243 F i g u r e 95. Growth o f o y s t e r s as a f u n c t i o n o f SIZE i n the two-stage c u l t u r e ( H| SIZE 1 fg| SIZE 2 E § SIZE 3 fx] SIZE 4)  EXP1  EXP  2  EXP1  EXP 3  EXP  2  EXP 4  EXP 3  § 20  I UJ  10,  PS  Q.  j EXP1  EXP  2  EXP 3  EXP  EXP1  EXP 2  EXP3  EXP4  4  4.0  rEXPi  EXP2  EXP3  EXP1  EXP 2  EXP 3  EXP4  EXP 4  244 Figure 96. The 'P versus I* curves as a function of temperature and n i t r a t e conditions.  TEMPERATURES \  >+C - -re  8  NON- NUTRi E.NT  cn  ' 0- A  UNITED.  x . -  CJ  o  CD .  rD \ u  o  O  &  A  •  •  O  Q  0 0  B  A 0.  0  +  A  •  +  9  0.0  0.1  T  r  1  o.z.  PAR  -1  1  1—  03  0.4  CU-WIN*11  r~—r  ~i  O  •  or  N U T R I E N T  U f V T f c D  \ cr. CJ  .  CD  \ CJ  O  o  0  CD«i'  Z3 €0  a.  0.0  A B  0.2.  a  © A  A A EI a  O  0.3  o.«f-  —i—  F i g u r e 97.  Comparison o f the s i m u l a t e d (PHYTO d u r i n g Experiment 5B: Run 1  ) versus a c t u a l  PHYTO MRX= 6 4 . 2 8 1 1 PHflV MRX= 5 4 . 1 7 5  TIME  (PHAV) p h y t o p l a n k t o n  stock  F i g u r e 98.  Comparison o f the s i m u l a t e d (PHYTO d u r i n g Experiment 5B: Run 2  ) versus a c t u a l  PHYTO MRX= 5 9 . 1 8 8 3 PHRV MRX= 5 4 . 1 7 5  TIME  (PHAV) p h y t o p l a n k t o n  stock  Figure 99. Comparison of the simulated (PHYTO during Experiment 5B: Run 3  ) versus actual (PHAV) phytoplankton stock  PHYTO MAX" 6 9 . 1 5 4 6 PHflV MRX= 5 4 . 1 7 5  TIME  NS  Figure 100. Comparison of the simulated (PHYT0——) versus actual (PHAV) phytoplankton stock during Experiment 5B: Run 4  PHYTO M f l X = . 7 9 . 0 8 8 6 PHflV MflX=. 5 4 . 1 7 5  TIME  N3 00  F i g u r e 101.  Comparison o f t h e s i m u l a t e d (PHYTO d u r i n g Experiment 5A: Run 1  ) v e r s u s a c t u a l (PHAV) p h y t o p l a n k t o n  PHYTO MflX= 6 2 . 8 2 1 4 PHRV MAX= 4 5 . 5 7 5  TIME  stock  F i g u r e 102.  Comparison o f the s i m u l a t e d ( P H Y T O — ) v e r s u s a c t u a l (PHAV) p h y t o p l a n k t o n d u r i n g Experiment 5A: Run 2  stock  PHYTO MflX= 4 4 . 6 4 8 7 PHRV MRX= 4 5 . 5 7 5  TIME  NJ Oi O  F i g u r e 103.  Comparison o f the s i m u l a t e d (PHYTO-—) v e r s u s a c t u a l (PHAV) p h y t o p l a n k t o n d u r i n g Experiment 5B: Run 1 w i t h a g r a z i n g term added on Day 27  PHYTO MAXPHRV MAX-  65.2397 54.175  TIME  stock  252  PLATE I I Artificial  c u l t c h i n the two-stage c u l t u r e experiments  253 BIBLIOGRAPHY A n d e r s e n , G.C. and Z e u t s c h e l , H.P. 1970* leiease of dissolved organic m a t t e r by m a r i n e p h y t o p l a n k t o n i n c o a s t a l and o f f s h o r e a r e a s o f t h e N. E. 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Winter,D.F., Baase. K and A n d e r s o n , G.C. 1975. The d y n a m i c s o f phytoplankton blooms i n P u g e t Sound, a f j o r d i n t h e N o r t h w e s t e r n United S t a t e s . , liath. Biosc. 29: 139-176..  260  A P E EN M X  VII *****  IX E 1  *  1.  D e s c r i p t i o n and p a r a m e t e r s used  EESCBIPT1CN 4**4*+;****  *  UNITS  *  variables  and  DEBIMTICls  *******  EXPERIMENT TIME  Z STN  ' £ AY  DEPTH STATION (INFL0H,OUTFLOR, S,M,E DEETHS) SUESTN SUBSTATION FLOW B A T E v VOLUME V FLUSHING BATE FB SINKING BATE SINK INCITENT Si SOLAB BABXATICN PHOTOSYNTHETICALLY IA B A V M L A I L E BADIAT * N PAH B U S I N G 0 2 E ABO INCUB*N PEBIOD PAS CUBING C 1 4 IABC INCUE'N PEBIOD EXTK EXTINCTION COEFF. EABAV PABZO  d e r i v a t i o n cf the i n t h i s study.  BAB AT DEPTB P A B AT DEPTH FOB 02 INCUE'N PEBIOD P A B Z C PAB AT DEPTH FOB C14 INCUB'N PEBIOD SALINITY SAL TEMPEEATUBE TEMP NET T E M P E E A T U B E TIME N {NITEATE] N03 NET [ N I T B A T E ] JSC3N [AMMONIA] K B 3 £ USEA ] OBEA £ PHOSPHATE] EC 4 £ SILICATE] SI03 [OXYGEN ] OXY NET £ OXYGEN ] OXYN OXYGEN S A T U S A T I O N SAT CHLA £CHLOEOPHYLL a] CfiLB £ CHLOEOPHYLL b] £CHLOEOPHYLL c] CHLC 1CAB0TEN0IDS] CI B A T I O C H L B TO C E L A EA B A T I O CHLC TO CHLA CA BATIO CT TO C E L A CTA B A T I O CHLB TO CHLC EC B A T I O C B L B TO CT ECT B A T I O CHLC TO CT CCT CAEBGN/CHLA :CHLA  METBE LITBE/DAY LITBE P E E DAY M/EAY LANCLEY/DAY (LY/DY) IANGLEY/DAY (LY/DY) L A N G L E Y / 4 HB (LY/INCUB) L A N G I E Y / 4 HE (LY/INCUB) P E B METBE  TIXED SAMPLING T (0830 H B EST) FIXED LOCATION' FIXED A B E A I IOC. MEASUBEC CAICUIATIE  v/V  M E l S USEE M E A SU BED 0.50*SB MEASUBEE MEASUBEC  . 0 4 * . GC88CHLA + .054 (CHLA**.6667) P A K E X P J-EXTK*Z) LANGLEY/MIN (. 5 * E A 1 0 / 2 4 G . ) * LAEGLEY/MIN E X P (-EXTK*Z) (LY/MIN) (.5*PABC/2 4 0 . ) * XAKGLEY/MIN E X P (-EXTK*Z) (LY/MIN) ME ASUBED PPT ( 0 / 0 0 ) MEASUBEC D E G B E E S C. (DEG-C) TEME ( Z ) - T E K E ( I ) DEG-C., MEASUBEC UM N 0 3 - N / L I T B E N03 ( I ) - N 0 3 (Z) UM N 0 3 - N / L I T B E MEASUBEC UM N H 3 - N / L I T B E MEASUBEC UM U E E A - N / I I T B E ME ASUSE D UM P G 4 - P / L I T B E MEASUBEC UM S I G 3 - S I / L I I B E ME ASUBED MG 0 2 / L I T B E OXY ( I ) - O X Y i ( Z ) MG 0 2 / L I T B E OXY/SOLUBIIITY PEBCENT (%) MEASUBEC UG C H L A / L I T B E MEASUBEC UG C E 1 E / L I T E E MEASUBEC UG C E L C / L I T B E MEASUBEC UG C T / L I T E E CBI-E/CHL fi DIMENSIONLESS C E L C / C Hi A DIMENSIONLESS CT/CHLA DIMENSIONLESS CELE/CELC DIHE NSIGNLESS C E I B / CT DIMENSIONLESS CEIC/ CT DIMENSIONLESS . CABEC S / C H L A DIMENSIONLESS  26 1  EGG I NO EES EBOD EXC PGOST FNOST BESST ASS EXCST EGGEY PCDY ALEE AG IX EH AC AEGO EEGO EPGO ESTPGO PMAX ETMAX PHYI'O EHAV P GIOWB GRAZE  GEOSS EBOD. (02) UG C/LITBE/HB NET EBOD, (02) UGC/LITBE/HE BESPIBATIGN UG C/LIT BE/HE NET PEOB. ,(C14) UG C/LITBE/HB EXUDATION (C14) UG C/LITBE/HB EGG (NORMALIZEE) UG C/UG CHLA/HE E NC(NO EM ALIZ ED) UG C/UG CHLA/HB BES (NOEMA LIZEB) UG C/UG CHLA/fiE PEOD (NOBM ALIZEE) UG C/UG CHLA/HB, EXC (NOB HA LIZ ED) UG C/UG C HI A/ HE DAILY EGO MG C/LITBE/DY DAILY EBOD MG C/LITBE/DY P.VS.I INIT.SLOPE UGC/UGCHLA/HE-LY/M P.VS.I INIT.SLOPE UGC/UGCHLA/HB-LY/M EBOD/PGO DIMENSIONS!ESS BES/PGO DIMENSIONSLESS EXC/EGO DIMENSIONSLESS ESTIMATED PGO DIMENSIONSLESS MAX. ASS UGC/UGCHLA/HB MAX., PMAX UGC/UGCHL A/HB SIMULATED CHLA UG CHIA/LITBE AVEBAGE CELA UG CELA/LITEE SIMULATED ASS UGC/UGCHLA/HB SIMULATED GBOWTH PEE BAY BATE SIMULATED BATE PEB DAY OF GBAZING  (UG 02/L/B £) /1.2 (EGO) - (BES) (UG 02/L/BE)/1.0 MEASUEEDCJ HB) M EASUBED (4 HE) EGC/CBLA ENG/CHLA EES/CELA EBOD/CHL fl EXC/CHLA PGO* (SI/EAIC) *4. EBOD* (SI/PAEC) *4. EGG/CHL A/P AE ZO EBOD/CEXA/E8BZC EECE/EGC BES/FGC EXC/FGG A P G 0 * E EG 0+11G 0 LSI ESTIMATE LSF ESTIMATE SIMULATION MEASOBED SIMULATION SI MUXATIC K SIMUIAIICN  26;  A1EENDIX 2.  VAE  4* *  *  D e s c r i p t i o n and d e r i v a t i o n o f a d d i t i o n a l v a r i a b l e s p e r t a i n i n g to h e r v i v c r e growth,  DESCB1ETIGN  «**4*4*  **<  *  1 « D WGTT WGTM WGTS NETL NETS NETD KETIT NET Hi! NETSS EE 11 PEEK IEBD PEE ET EEiWM EEEWS GB KM GEHS MSBATIO ESUEV SIZE  LENGTH WIDTH DEPTH TOTAL WEIGHT MEAT KEIGHT SfcILL WEIGHT NET LENGTH NET WIDTH NET DEPTH NET TOTAL WEIGHT NET MEAT WEIGHT NET SHELL WEIGHT % INCB. LENGTH % INCB. HIDTH 31 INCB. , DEPTH % ISC8. WGTT % INCB. KGTM % INCB. WGTS NETJsM/WEEK N ETWS/fi f EK NETH.fi: NET8S # SU1V1VCBS HERBIVOBE SIZE  BESS  HEREIVOEE DENSITY  STKUP STROPS CXYUP OXYDPR  UPTAKE CCNC. OI PHYTOELANKTON UPTAKE BATE GE PHYTOPLANKTON UPTAKE CONC. OF OXYGEN UPTAKE BATE OE OXYGEN  UNITS  * * * * * * <  Cfl CIS  CM GRAMS GRAMS GRAMS ce  CM CM GRAMS GJ3AHS GBAMS  G/ZGO/WK G/ZOC/WK DIME NSIC NLESS NUMEEB N/A N/A UG CHI A/LIT EE UG CHLA/DAY MG 02/IITBE MG 02/IITBE  *  DERIVATION  * * * * * * *  * * * * * * * * * * * * *  EEASUSED MEASURED ME ASU.RI D MEASURED EEASUEED MEASURED I (f)-L U) s<f)-i<i) r (f)-E'ti) WGTT (f) -WGTT { i) WGTM (f) -SGTE'(i) WGTS (f) -WGTS (i) KETI/Ifi) NETW/W<i) NETD/D (i) NETM/WGTT (i) NETKM/fiGTM(i) NETWS/WGTS(i) CALCULATED CALCULATED CALCULATED Factor  with U classes <1=£tallest;4=1erg est) F a c t o r with 3 classes <1=lowest;3=highest) CHIA <I) -CHLA (C) STKUP*IB OXY (1) -CXY (C) CXYUP*IB  

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