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Preliminary exploration of estuarine ecosystem structure at low trophic levels with a controlled microcosm Wu, Yong 1985

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PRELIMINARY EXPLORATION OF ESTUARINE ECOSYSTEM STRUCTURE AT LOW  TROPHIC LEVELS WITH  A CONTROLLED MICROCOSM  by Yong B.Sc,  Wu  Xiamen U n i v e r s i t y ,  Xiamen, C h i n a ,  A THESIS SUBMITTED IN PARTIAL THE REQUIREMENTS FOR MASTER OF  FULFILLMENT  THE DEGREE OF  SCIENCE  in THE FACULTY  OF GRADUATE  STUDIES  (Department o f Oceanography)  We  accept to  this  thesis  as  the required  THE UNIVERSITY  standard  OF BRITISH COLUMBIA  December, (c)  conforming  1985  Yong Wu,  1985  1982  OF  In p r e s e n t i n g requirements  this thesis f o r an  of  British  it  freely available  Columbia, I agree that for reference  by  understood that for  h i s or  be  her  copying or  f i n a n c i a l gain  shall  3 March,  1986  Library  s h a l l make  and  study.  I  publication be  the  of  further this  Columbia  thesis  head o f  this  my  It is thesis  a l l o w e d w i t h o u t my  Oceanography  The U n i v e r s i t y o f B r i t i s h 1956 Main Mall V a n c o u v e r , Canada V6T 1Y3  Date  University  representatives.  not  the  the  g r a n t e d by  permission.  Department o f  the  f o r extensive copying of  s c h o l a r l y p u r p o s e s may  department or  f u l f i l m e n t of  advanced degree a t  agree t h a t p e r m i s s i o n for  in partial  written  Abstract  In were  a  laboratory  created  circulation of  the  with  simulate mixing  freshwater  to  the  were  chosen  of  the  sea water  gradients  the  pattern  of The  the  different  gradients  and d i f f e r e n t stage  I  salinity  freshwater  biota  Most  of  them  The  autotrophic as  by  the  salinity  homogenous  could  autotrophic  in  autotrophic  among  with  component  The  i n  ecosystem the  the  and  difference  different  data  through  salinity  fresh  develop  were t e s t e d on  two  showed  the  gradient.  the mixing  water  a  that  the salinity  during  component  could  process.  no  longer  as an  organic  of phytoplankton  growth  but served  c o n d i t i o n i n which i n  the  controlled flasks, was  i n  origin.  provided  concurrently  i nthe  interaction  reflected  the  with  The i n h i b i t i o n  gradient  condition  was  reflected  inhibited  component  contributor.  bacteria  the  not pass  o r were  trophic levels  phytoplankton  gradients.  parameters  n u t r i e n t s , b a c t e r i a , and  experiments  ecosystem  could  died  a  were  freshwater  different  levels  behaviour  i n relation  c o n d i t i o n s . The  interaction  between  The  i n v e s t i g a t e d . Seven  phytoplankton,  conditions  substrate  were  gradients  i n estuarine  ecosystem components  trophic  environmental  function  occurring  and f r e s h w a t e r .  environmental  different  nanozooflagellates.  In  process  of salinity  as i n d i c a t o r s o f t h e d i f f e r e n t  or  development  two k i n d s  the  and seawater  salinity  ecosystem among  to  microcosm,  separated  ecosystem.  the In a  t h e development of from  heterotrophic  i i i bacteria  over  In from  stage  that  growth  time.  of  ecosystem  had a d i f f e r e n t  fresh  With  increasing  water.  of phytoplankton could  resulted  in  provided  the  before a  I I , a seawater  a  delay first  before  could,  on  growth,  This  o f t h e maximum p h y t o p l a n k t o n b i o m a s s  and  of  time  phytoplankton  component  the salinity may  play  developed.  phytoplankton  gradient.  the  major  estuarine  autotrophic  components  i n both  on  o f t h e components  cause  a  great  change  developmental different which the  experiments  function  were  the  of  of  idea  development  systems  The  their  autotrophic  behavior  explained with  and  could  Different  bacteria  a conceptual  states  in  in  diagram  o f an ecosystem  bacteria  of  effect  changes  structure.  phytoplankton  and  i n their  can have a s t r o n g  i n t h e system;  phytoplankton  respond  i n primary production i n a  ecosystem.  o f energy  formed  phytoplankton  Thus t h e s e a w a t e r  i n t h e whole system  patterns  summarizes  Seawater  and q u i c k l y  role  based  the rest  forbacterial  components. Thus b a c t e r i a  t h e o t h e r hand, a c t i v e l y  on  salinity, the  a t low s a l i n i t i e s .  period  be l i m i t e d  t h e bloom o f a u t o t r o p h i c  peak  an  responce  and  ecosystem  dynamics. Temporal our if  experiments a  body  conceptual distribution have  development  often  estuaries.  of  may  be e x t r a p o l a t e d  water  model of  i s many  been  p a t t e r n s o f the ecosystem  i s  moving  seaward  presented biologically  reported  i n  into  to  spatial  components i n distributions  i n an e s t u a r y . explain  important many f i e l d  the  Thus a spatial  components  which  investigations of  iv TABLE OF  CONTENTS  Page ABSTRACT  i i  TABLE OF CONTENTS  iv  L I S T OF TABLES  V  L I S T OF FIGURES  vi  ACKNOWLEDGEMENTS  ix  1.  INTRODUCTION  2.  EXPERIMENTAL DESIGN  10  2.1 PRECONSIDERATION  10  2.2 DESIGN OF THE EXPERIMENT  14  2.3 CONDITION OF EXPERIMENT  18  METHODS OF ANALYSIS OF SAMPLES  22  3.1 NUTRIENTS  22  3.  3.2 TOTAL BACTERIA NUMBERS, BIOVOLUMES  4.  1  AND  BIOMASS  23  3.3 CALCULATION OF SALINITY GRADIENT  25  3.4 MICROFLAGELLATES AND PHYTOPLANKTON  25  3.5 STANDING STOCK OF PHYTOPLANKTON  26  RESULTS  28  4.1 EXPERIMENT STAGE I  28  4.11 INCREASING SALINITY GRADIENT  28  4.12 DECREASING SALINITY GRADIENT  48  4.2 EXPERIMENT STAGE I I  60  4.21 INCREASING SALINTIY GRADIENT  60  4.22 DECREASING SALINITY GRADIENT  69  5.  DISCUSSION  6.  LITERATURE CITED  84 109  V List  of Tables  Table page 1. The p r o c e d u r e s o f t h e e x p e r i m e n t -in t h e two s t a g e s . 17 2.  The s a l i n i t y o f i n i t i a l times f o rthe l a b o r a t o r y  waters c o l l e c t e d a t experiments  different 22  3.  C o r r e l a t i o n m a t r i x o b t a i n e d f r o m t h e 7 v a r i a b l e s X 20 observations i n experiment stage I, part 1 during J u l y , 1985, between t i m e ( d a y ) , F l u o r e s c e n c e ( f l u o ) , N0~3, NH+4, F l a g e l l a t e s ( f i g ) , B a c t e r i a (bac), and S a l i n i t y . P r o b a b i l i t y : * p<0.1, ** p < 0.05, *** p < 0.02, **** p <0.01 41  4.  C o r r e l a t i o n m a t r i x o b t a i n e d f r o m t h e 7 v a r i a b l e s X 20 observations i n experiment stage I, part 2 during J u l y , 1985, between t i m e ( d a y ) , F l u o r e s c e n c e ( f l u o ) , NO~3, NH 4, F l a g e l l a t e s ( f i g ) , B a c t e r i a (bac), and S a l i n i t y . P r o b a b i l i t y : * p<0.1, ** p < 0.05, *** p < 0.02, **** p <0.01 41 +  5.  C o r r e l a t i o n m a t r i x o b t a i n e d f r o m t h e 6 v a r i a b l e s X 12 o b s e r v a t i o n s i n experiment stage I I , p a r t 1 during Aug., 1985, between t i m e ( d a y ) , F l u o r e s c e n c e ( f l u o ) , N0~3, F l a g e l l a t e s ( f i g ) , B a c t e r i a ( b a c ) , a n d s a l i n i t y . P r o b a b i l i t y : * p<0.1, ** p < 0.05, * * * p < 0.02, **** p <0.01 68  6.  C o r r e l a t i o n m a t r i x o b t a i n e d f r o m t h e 6 v a r i a b l e s X 12 o b s e r v a t i o n s i n experiment stage I I , p a r t 2 during Aug., 1985. between t i m e ( d a y ) , F l u o r e s c e n c e ( f l u o ) , NO~3, F l a g e l l a t e s ( f i g ) , B a c t e r i a ( b a c ) , a n d S a l i n i t y , p r o b a b i l i t y : * p<0.1, ** p < 0.05, *** p < 0.02, **** p <0.01 68  vi L I S T OF F I G U R E S  1.  Flow diagram f o r t h e upper l a y e r box o f water d i m e n s i o n a l model o f e s t u a r i n e c i r c u l a t i o n  2.  Sampling s i t e s f o r laboratory experiments and surface s a l i n i t y d i s t r i b u t i o n i n Fraser estuary and plume  17  3.  Design  20  4.  The t e m p o r a l s a l i n i t y p a t t e r n i n t h e e x p e r i m e n t s t a g e I, I I p a r t 1 w i t h an i n c r e a s i n g s a l i n i t y g r a d i e n t 29  5a.  The development p a t t e r n o f a u t o t r o p h i c component and nutrient i n t h e experiment stage I , part 1 during July, 1985 30  5b.  The development p a t t e r n o f NH 4 i n t h e e x p e r i m e n t s t a g e I , p a r t 1 d u r i n g J u l y , 1985  o f the experiments with  three  i n a two 11  units  +  31  6a.  The development pattern of autotrophic nutrient i n the experiment stage I , part F e b r u a r y , 1985  component and 1 during 32  6b.  The development p a t t e r n o f NH 4 i n t h e e x p e r i m e n t s t a g e I , p a r t 1 d u r i n g F e b r u a r y , 1985  +  33  7.  The development p a t t e r n f o r b o t h h e t e r o t r o p h i c b a c t e r i a and n a n o z o o f l a g e l l a t e s i n e x p e r i m e n t s t a g e I p a r t 1 d u r i n g J u l y , 1985 36  8.  Growth o f f r e s h w a t e r b a c t e r i a on p l a t e different salinity concentrations  9.  media a t  The t e m p o r a l s a l i n i t y p a t t e r n i n e x p e r i m e n t s t a g e I I , part 2.with a decreasing s a l i n i t y gradient  .36 I & 43  10a.  The development p a t t e r n o f a u t o t r o p h i c component and n u t r i e n t s i n S t a g e I , p a r t 2. w i t h a d e c r e a s i n g s a l i n i t y g r a d i e n t i n J u l y , 1985 44  10b.  The development p a t t e r n o f NH 4 c o n c e n t r a t i o n i n Stage I , part 2 with a decreasing s a l i n i t y gradient i n J u l y 1985 45  11a.  The development p a t t e r n o f a u t o t r o p h i c component and n u t r i e n t s i n Stage I , p a r t 2 with a decreasing salinity g r a d i e n t i n F e b r u a r y , 1985 46  +  Vll lib.  +  The d e v e l o p m e n t p a t t e r n o f NH 4 c o n c e n t r a t i o n i n Stage I, P a r t 2 with a d e c r e a s i n g s a l i n i t y g r a d i e n t i n F e b r u a r y , 1985 47  12. The d e v e l o p m e n t p a t t e r n o f b a c t e r i a a n d n a n o z o o f l a g e l l a t e s i n Stage I , p a r t 2 with decreasing s a l i n i t y gradient  a 49  13. The d e v e l o p m e n t p a t t e r n o f a u t o t r o p h i c component, n u t r i e n t and b a c t e r i a i n a c o n t r o l l e d s y s t e m w i t h o r i g i n a l seawater d u r i n g J u l y , 1985  53  14. The d e v e l o p m e n t p a t t e r n o f a u t o t r o p h i c component n u t r i e n t and b a c t e r i a i n a c o n t r o l l e d s y s t e m w i t h o r i g i n a l f r e s h w a t e r d u r i n g J u l y , 1985  54  15. The d e v e l o p m e n t p a t t e r n o f p h y t o p l a n k t o n and c o n c e n t r a t i o n o f n u t r i e n t s i n s t a g e I I p a r t 1 on i n c r e a s i n g s a l i n i t y g r a d i e n t d u r i n g Aug. 1985  61  16a. The d e v e l o p m e n t p a t t e r n o f a u t o t r o p h i c component and c o n c e n t r a t i o n o f n u t r i e n t s i n s t a g e I I p a r t 1 on a n i n c r e a s i n g s a l i n i t y g r a d i e n t d u r i n g A p r i l , 1985 62 16b.  +  The d e v e l o p m e n t p a t t e r n o f NH 4 i n s t a g e I I , p a r t 1 on an i n c r e a s i n g s a l i n i t y g r a d i e n t d u r i n g A p r i l , 1985  17. The d e v e l o p m e n t p a t t e r n o f b a c t e r i a a n d n a n o z o o f l a g e l l a t e s i n stage I I , p a r t 1 with s a l i n i t y g r a d i e n t d u r i n g Aug., 1985  63  increasing 65  18. D e v e l o p m e n t o f a u t o t r o p h i c component i n s u b s a m p l e culture s e r i e s a t temporally constant s a l i n i t y i n s t a g e I I , p a r t 1 d u r i n g Aug., 1985  66  19. T h e d e v e l o p m e n t p a t t e r n o f a u t o t r o p h i c component a n d c o n c e n t r a t i o n o f n u t r i e n t s i n s t a g e I I , p a r t 2 on a d e c r e a s i n g s a l i n i t y g r a d i e n t d u r i n g Aug. 1985 70 20a. The d e v e l o p m e n t p a t t e r n o f a u t o t r o p h i c component and c o n c e n t r a t i o n o f n u t r i e n t s i n s t a g e I I , p a r t 2 on a d e c r e a s i n g s a l i n i t y g r a d i e n t d u r i n g A p r i l , 1985 71 +  20b. The d e v e l o p m e n t p a t t e r n o f NH 4 i n s t a g e I I , p a r t 2 on a d e c r e a s i n g s a l i n i t y g r a d i e n t d u r i n g 1985 21. The d e v e l o p m e n t p a t t e r n o f b a c t e r i a a n d n a n o z o o f l a g e l l a t e s i n s t a g e I I , p a r t 2 on a s a l i n i t y g r a d i e n t d u r i n g Aug., 1985  April, 72  decreasing 75  viii 22. D e v e l o p m e n t o f a u t o t r o p h i c component i n a s u b s a m p l e culture s e r i e s at temporally constant s a l i n i t y in s t a g e I I , p a r t 1 d u r i n g Aug., 1985  76  23a.  The development p a t t e r n o f a u t o t r o p h i c component i n a c o n t r o l l e d system with o r i g i n a l seawater d u r i n g Aug. 1985 82  23b.  The development p a t t e r n o f a u t o t r o p h i c component i n a c o n t r o l l e d system with o r i g i n a l freshwater during Aug. 1985 83  24.  The dynamics energy f i e l d  25.  of ecosystem  energetic  The arrangement o f e c o s y s t e m physical force field.  state  i nthe 88  components  i n an  estuarine 105  ix Acknowledgement This t h e s i s would not have been p o s s i b l e w i t h o u t the h e l p o f many people. I am v e r y g r a t e f u l t o my t h e s i s s u p e r v i s o r , D r . T . R. Parsons, for his encouragement, i n s p i r a t i o n i n the i n i t i a t i o n and the freedom of exploration throughout the study. It is he who directed me in this f i e l d and I think his influence in t h i n k i n g o f e c o s y s t e m s w i l l b e w i t h me f o r m a n y y e a r s t o c o m e . I am grateful to Dr. P. J . Harrison for his constructive suggestions, helpful discussion as well as the k i n d n e s s he showed to me d u r i n g the p a s t two y e a r s , and a l s o f o r p r o v i d i n g the use o f h i s laboratory. Special thanks are due to Dr. A. G. Lewis for his very helpful discussion and comments at the beginning of my laboratory w o r k . Thanks are a l s o due t o D r . F . J . R. T a y l o r , f o r his comments and allowing me to use h i s l a b o r a t o r y . Here, thanks are extended to Miss J u d y A c r e m a n f o r h e r h e l p i n my identification of species. Much appreciation is extended to Dr. Carol L a l l i for her w o r k i n i m p r o v i n g my m a n u s c r i p t . I am g r a t e f u l t o D r . R . J . A n d e r s e n , w h o s a t o n my c o m m i t t e e a n d f o r a p p r o v a l o f my p r o p o s a l a n d r e a d i n g my t h e s i s . I like to thank D r . P . H . LeBlond and D r . R.W. B u r l i n g for their h e l p f u l d i s c u s s i o n on the e s t u a r i n e c i r c u l a t i o n . My a p p r e c i a t i o n i s a l s o g i v e n t o many f e l l o w s , f a c u l t y and secretaries at the Department of Oceanography for t h e i r kindness a n d t h e h e l p t h e y h a v e g i v e n me w h e n e v e r I n e e d e d i t . Special thanks to Miss Heather Dovey for her friendship and much assistance throughout the study and her a s s i s t a n c e in typing this manuscrip and in drawing the f i g u r e s . Special thanks to Peter Thompson f o r h i s k i n d a s s i s t a n c e d u r i n g my u s e of Dr. Harrison's laboratory equipment and the helpful discussion during my e x p e r i m e n t . T h a n k s a r e d u e t o M . P . S t o r m for his assistance i n m e a s u r i n g s a l i n i t i e s f o r my e x p e r i m e n t s . Many thanks to my fellow graduate students, for their friendship and g e n e r a l h e l p w h i c h g a v e me a g o o d e n v i r o n m e n t in the past two years, and many thanks to students i n Dr. P . J . Harrison's group for t h e i r general help. To d e t a i l a l l of the support I have r e c e i v e d i s i m p o s s i b l e . I would l i k e t o t h a n k a l l my f r i e n d s i n C a n a d a a n d i n C h i n a for their continuous encouragement, care and h e l p which d i r e c t l y and i n d i r e c t l y a i d e d me i n m y t w o y e a r s o f study. I w i l l always be grateful to my p a r e n t s a n d my w i f e for their constant moral support and the c o n f i d e n c e they p l a c e d i n me; t o them I d e d i c a t e t h i s thesis. Finally, I a c k n o w l e d g e t h a t I was s u p p o r t e d by t h e N a t i o n a l Scholarship of The P e o p l e s ' R e p u b l i c o f C h i n a and an award from the International Development Research Centre of Canada. hard  1 1.  Estuarine and  ecosystems  marine  systems  relatively sea  very  water  space  mix  which  Pritchard typical  are  and,  drives  (1955)  a  as  complex.  together  being  Introduction  such,  Within  to  the various  partial  typical  circulation  spatial  and  highly  associated  with  ecological conditions are related  transition a  there  must  f o r both  convenient  way  of  for  identifying  the  the  ecosystems  along  using  causes  a  landward  components subject the  flows  to  which  the  are  on  Therefore,  t h e most  distinctive  Moreover,  This  i n  along  of  salinity  )  water  gradient  most o f t h e  Obviously,  underneath,  of  extremes o f  on t h e s u r f a c e  both  provides  dynamic  filter.  this  behaviour  both  ( Kennedy, 1984 this  gradient.  association nature  many  which are  to this  zone from  estuary  suspended process  has  and temporal  flow of sea water  the mixing  transition.  filter  react  into  This  spatial  29°/oo).  a biological  t h e most  parameters  the conservative  to  as  water  systems.  transition  functions  patterns.  conditional gradient  this  0°/oo  fresh  a  dynamic  (  parameters  be  source  salinity  biological  important  over  with  a  gradient.  composed  ,  and  f r e s h water and  which  and  Therefore  dynamic  patterns,  estuary  salinity  water  a density difference  biologically of  fresh  estuarine circulation  mixing  ecologically  between  an e s t u a r y  four  pattern  temporal  are  produce  classified  a  transition  when  l a y e r and  the ecosystem bodies  will  be  on t h e c o n d i t i o n a l g r a d i e n t  along  one  broad  may  address  a  very  2 question what  of  ways  does  functional the  what  environment  one  the  component  result  in  a  change  from  while  more  from as  in  ecological  (Lauff,  both  marine  changes  or  important and  function  ecosystem.  The  and  the than  fresh  water  another  of  This  energy  and  and  quantities  e s t u a r y may  It is  systems undergo  original  e v e n w i t h o u t man  be  interesting  the  of  kind  through.  either true  areas  so  hybrid estuarine  added p o l l u t a n t s .  a  the  a hybrid  resulting  other  is  from  circulation  of  quality  of  a converter,  pass  estuary receives.  is  this  into they  exported  dynamic  type  t o form  as  in  limiting  one and  distribution  This  nursery  from  processes  ) when  a  p e r m i t t i n g or  changed  different  components  productive  sewage  be  the  by  ecosystems  may  gradient, or  g r a d i e n t a c t as  mixing  t o DOC  the  those  1967).  species  may  POC  environmental are  do  salinity  components  very different  as  different that  two  How  (such  structure  the  ecosystem  This gradient f i l t e r  system's  would  either  ecological  couple  component  of  on  of  of  conditional  to another?  ecosystem?  effect  this  filter  passage  patterns  i s the  water  made  extreme systems bodies  enrichment  E s t u a r i e s have been  used  f o r many e c o n o m i c a l l y i m p o r t a n t  fish  reflection  of  their  high  primary  productivity. An the  increasing structure,  stimulated knowledge 1967),  the  comparison  a  demand function,  number  of  the  of  the  a comprehensive  and  dynamics  conferences  natural  interactions of  for  and  characteristics of  these  different  of  estuaries  symposia of  systems  processes  understanding  has  summarizing  estuaries  (Lauff,  (Wiley, 1976), in  these  of  the  estuarine  3 systems 1984  ( Kennedy, 1982),  ),  the  1979).  dynamics  Microbial  estuarine from  comprehensive numerous  understanding  from  to  estuaries be  with  have  helpful  fully to  Several  1978; found  (or  and  low  Goulder  Wright, stock the  have  et  strong  1983) often  aspects. which  obtaining  the  about  our  proving  the  system  which  at whole  better  a  ecosystem.  i t is  Studies  look  a  increase  treat  processes  p i c t u r e of  e c o l o g i c a l s t r u c t u r e of  perspective found  an  Foester,  Seliger  zone  our  along (  and  in  Recently,  unique  understanding  specific  attention  included  which  systems  basic  the  involved.  studies  gradient  in  understand  focus  parameters  overall  of  (1977).  this  Cronin,  aspects  were  have appeared  dynamic  &  preliminary and  about  ( Kennedy,  (Neilson as  Rheinheimer  whole ecosystems or  structure  To  (1973)  knowledge  dealing  estuaries  nutrients  received  by  arevery  of  physiology  Colwell  an  more  dynamic  have  construct  as  and  about e s t u a r i e s and  studies  may  and  estuaries  difficult  of  also  functions  chemical  discussion  papers  However,  of  ecology  ecology  Stevenson  the  a  a l . ,  1981;  and  at  the  (Parsons,  1983;  activity  of  the an  we  estuary.  most e c o l o g i c a l or  salinity  Palumbo and  Ferguson,  zone  the  (Allbright,  mouth of 1969;  of  1978).  at  al.1982;  maximization  appears  gradient)  et  pattern  Wright,  regions)  Joint  s c a l e w i t h i n an  transition  Helder,  heterotrophic salinity  the  spatial  estuarine 1976;  1981;  at  estuaries  Morris  turbidity  maximum  1977,1983a,b,;  Valdes  et  estuary  (or  1984;  Bent  al.1980,1981;  phytoplankton  Cadee 1978,  (1978)  the  standing sea  Stockner  end et  4  al.,  1979;  Harrison,  ecological  parameters  distribution 1979;  Peterson,  understand  the  through  the  kessler,  1984),  Ecosystem between  the  terms  covariance  is  system  position  field of  trace (detritus) included  estuarine  of  1981;  phytoplankton,  Second, spatial  within  this  and d y n a m i c s o f  knowledge  of  both  mechanisms. are in  organic  two  the  major  water  and  organisms  flagellates,  levels masses:  nutrients,  material  living  bacteria,  the  components  inorganic  2).  components  levels.  and m o d i f y  requires  there  ions,  materials,and  the inherent  flow determine the  the structure  contained  dissolved  First,  energy  ecological  flow,  including  metals,  Cheng,  lacking.  among t r o p h i c  ecological  compartments  substances;  and  functions  p h y s i c a l mechanisms and  ecological  and  either  o f r e a c t i o n s w i t h i n and  may a r r a n g e  understanding  ecosystem  an  phenomena  among t h o s e e c o s y s t e m  components  temporal  is still  and e c o s y s t e m .  those  and  In  these  the r e s u l t  cycling  forcing  estuarine  1984;  have been made t o  (Loern  picture  material  good  Cadee,  P e t e r s o n a t a l . , 1984) o r s i m u l a t i o n  but a c l e a r  physical  A  dynamic  ( M a r g a l e f , 1967; C o o p e r e t a l . , 1973; S p i e s &  the  field.  a l l other  type o f s p a t i a l  attempts  behind  interrelationships  of  Almost  a l . , 1982;  modeling  1985;  physical  of  et  f a r , some  structure  (intrinsic) in  So  mechanisms  laboratory  Parsons,  show t h i s  Sharp  mathematical  Parsonsand  1984).  1982; Kemp e t a l . , 1982; N e i l s o n e t a l . ,  1975;  1985).  Peterson,  also  (Ducklow,  Carpenter,  in  1983;  of 1).  biological particulate  which  mainly  ciliates  and  5 macrozooplankton  (the l a t t e r  ).  about  The  concept  ecological new  which  flagellates 1963, in  in  Parsons  the  than  that  that  of  of  the  (DOC+POC)  is  1979;  in  et  Bacteria  (Rhee,  other  nutrients  1979;  Vaccaro  biomass  can  1985).  of  In  than large  e s t u a r i e s , the  Both  the  ( Neilson et  and  bacterial  a l . , 1978;  bacteria  in  the  fauna  Bratbak,  total 1984,  as  compete  p h o s p h o r u s and Saks,  g r o w t h r a t e s and  of bacteria  (Biddanda,  i n estuarine  established  (Wright,  n e t p l a n k t o n and l a r g e phytoplankton  recycling  K i n g e t a l . , 1980). system  can  high quantity  equal r o l e with nutrient  of inorganic  Fredrickson,1977;  h a s been f i r m l y  p l a y an  they  inorganic  position  as i m p o r t a n t  et  scavengers  1975),  significantly  Sieburth  (  less  i s a very  i n t h e ocean  for  1975,1981;  a  flow  ecosystem.  Parker,  only recently  energy  there  are e f f i c i e n t  a l . , 1969).  Bacteria  i s 5 t i m e s more  u s e r s o f DOC and POC  They a r e a t l e a s t  organisms.  o r g a n i c carbon(POC)  have been known a s  Williams,1981).But  ecosystems  as  1984; W i l l i a m , 1975;  (Brown,  reach  early  a l . , 1980; W r i g h t ,  phytoplankton  et  As  b i o m a s s and 10 t i m e s  than t h a t  1972;  with  as a r e s u l t o f  1971,1975).  So  of these  o f b a c t e r i a and  2 X lO^-^ g w h i c h  carbon.  higher  role  particulate  aquatic  long time. B a c t e r i a  successfully  terms  the  Billen  nutrients  1985;  organic  much  Turner,1978). a  out that  phytoplankton  dissolved pool  for  ecosystems(Sorokin,  amounts t o a b o u t  bioenergy  al.,  has changed d r a m a t i c a l l y  comfirms t h e important  pointed  ocean  i n our experiments  t h e s t r u c t u r e and i n t e r a c t i o n s  compartments  evidence  are not included  may a p p r o a c h  in  ( S o r o k i n , 1977;  The b i o m a s s o f t h e that  of the  entire  1985; Fuhrman & Azam,1982 N e w e l l , 1 9 8 1 ;  6 Watson,  1979).  faster of  than t h a t  energy  The  Since  and  very  Because  direct  1981). to  90%  difference  can  the  Goldman,  opens  & a  Following  (Eppely  &  1985;  form.  A  "the the  state  the  ecosystem and  (Thomas,  1971;  Bacteria  use  flow  by  of  a  very  Larsson  and  the  detritus  to  (Cobeney, 1982;  (Williams,  1984;  Fenchel,  is  are This  evidence  that  of  bacteria Caron  and  1982a, b,  c,;  grazing  recycled  important  80  1979).  main g r a z e r s  process back  to  in  the  component  thus  established.  e n e r g y pathway, t h e d i s t r i b u t i o n s t a r t i n g from p h o t o s y n t h e s i s  has  been  is  subject estimated  immediately  for that  released  & H a g s t r o m , 1982,-Smith and  energy  are  that probably  Peterson,  interesting  It  nutrients.  zone o f p e l a g i c w a t e r  loop",  energy flow  1978).  (50%-70%)  A n d e r s e n e t a l . 1985;  this  flux  apparently  abundance o f  very  phytoplankton  this  in bacteria  s u b s t a n c e s t o be  microbial  the  an  are  and  reconstruction.  substances  by  that  (Marita,  nutrients  and  Sieburth,  new  became  fixed  great  Haas & Weebs, 1 9 7 9 ) . The  discovery of  i t follows  demonstrated  euphotic  explained  Davis  Caron,  and  energy  i n the  (Azam e t a l . , 1983;  ecosystem,  carbon  have  pathway f o r o r g a n i c  inorganic  study  be  1985;  Goldman  inorganic  microflagellates  sea  b a c t e r i a i s much  i n r e m i n e r a l i z a t i o n of  of  regeneration  phagotrophic  of  a m a j o r mode f o r r e m i n e r a l i z a t i o n  nutrients  by  t o be  efficiencies  a gap  excretion  i s not  of  likely  conversion  However many s t u d i e s  supplied  in  is  leaves  this  time  of macroorganism biomass,  high  1977)  ,  turnover  material  (Calow,  low  the  Wiebe  source both from d i s s o l v e d create Cole,  a  large  e t a l . 1982;  further 50%  of  as  DOM  1976). organic  biomass f l u x Lancelor,  in  and  1979).  7 Through  this  serves  to  feeding  repackage  zooplankton  recycled and  "microbial  back  Goldman,  in  this  l o o p may  and  ( F a u s t , 1976; play  Because have  to  these  energy the  two  and  may  difference  of  patterns  reveal  functional  the  realistic  different  1977).  of  each  water  column.  of  in this  f o r energy pathways  et a l . ,  The  1983;  flow exist, The  dynamic  response  organic microbial  coexistence.  they  must  distributions of the  time  i n terms  of  differences  these  environmental  differences The  of  ecosystem  behaviour of  to the  functional  from  the  found  ecosystem.  flow.  components  and  So  i s a response  their  and  evolution  their in  the  growth  adaptation  commensalism, c o m p e t i t i o n and p r e d a t i o n of  t h e two  the environment.  estimates  pathways,  made  to  (Caron  have been  & Conrell,  interrelationship  response  i t is  activity  result  interacting,  of  bacterial  pathways of energy  which  capability,  (Linley  Faust  two  part  filter  minerialized)  nutrients  their  to  support high populations  the d i f f e r e n t  these  (i.e.  and  biomass  estuaries  of  reflect  of  1979)  inorganic  functions  components  accessible  investigations  environmental conditions.  significance  some  two  the b a c t e r i a l  both  in different  conditions  will  Webb,  a more i m p o r t a n t r o l e  different  the  for  of  cells  substances  Many  high rates  into  and  shown t h a t  system  compounds  (Haas  inorganic  have  bacteria  energy  1985).  Wright,1984) of  to  loop", part  of  carbon  provided that  the components of  This provides the  pathways and  It i s possible flow  through  their  t o make the  simultaneous measurements the  basic  standing stock i n the concept  i n our  two are same  experiment.  8 The  questions  addressed  essential  processes  1)  is  What  the  ecosystem  bacteria  and state  dilution  process?  What  of  are  compartments  the  the  when  dynamic  they  go  3)  components  temporal  the  o f some f r e s h w a t e r and  salinity  salinity  estuarine  rate?  a l l the  some b a s i c and  ecosystems:  and r e s p o n s e  to  constant d i l u t i o n Do  a r e about  components e . g . , p h y t o p l a n k t o n ,  nutrients  dynamic  2)  i n estuarine behaviour  seawater  here  gradient  environment  interactions through  gradient  in  as  among  the  they  and t o t h e  i n terms  of the  these  the s a l i n i t y  behave  flagellates,  major  gradient  at a  same manner on t h e  did  in  the  original  environment? 4)  What  is  assemblages 5)  How  in  can  components (in  the  both  the  all  attempt  other  etc.  These  on t h e g r o w t h o f p h y t o p l a n k t o n  and s e a w a t e r ?  in  growth  be e x t r a p o l a t e d  patterns  of d i f f e r e n t  for spatial  significance  an e s t u a r i n e c i r c u l a t i o n p a t t e r n ) ? answer  associated  gradient;  microcosm  f r e s h water  time  to  light  effect  difference  over  considering To  salinity  t h e s e q u e s t i o n s , I have t o e l i m i n a t e  factors  temperature  artificial experiments  i n nature such a s : t h e h o r i z o n t a l difference;  sediment  influences,  c o n d i t i o n s c a n be a c h i e v e d i n l a b o r a t o r y and  an  apparatus  was  constructed  to  achieve the objective. The  purpose  investigation primary  of  trophic  of the levels  this  work  was  time e v o l u t i o n along  a  to  make  a  preliminary  o f growth p a t t e r n s temporal  salinity  o f two  gradient.  9 Estuarine so  ecosystems  many  features  experiments this  on  the  variety that an  low  subeffect  physiological  different  by  The  understand growth  to  of  the  of  some  of  pattern to  ecosystems  which water  high  i f the  of  high  is  In o r d e r  to  observe  other  unknown  to  r a t e and are  only  to  time  simulated  microcosm;  described  and  a  or the  i n Chapter by  growth  i n Chapter I I I .  interacting  during  light  effect  gradients  these  and  of  turbidity  s t r u c t u r e produced  of  shown  segment  become one  or  are  described  structure  i n a wide  n u t r i e n t - r i c h deep water  design the  on  identify  work has  t i m e w i t h i n one  arrangement  happened  found  and  to d i l u t i o n  saline  and  the  masses.  area  but,  to  salinity  question:  salinity  purpose  the  low  salinity  over  b i o t a are  immediate  a d u p l i c a t i o n of  i n order  turbidity  (due  laboratory  imitate nature  r e a l mechanism.  temporal  water  be  turbidity  the  the  (due  development  pertaining  different  the  at the  that  e v o l u t i o n p a t t e r n and  differences The  ask  does  entrainment fresh  experimental II.  may  physical effects  constant  dilution  masks  to  situation  high  then  effects  and two  not,  not  example, p r e v i o u s  the  So we  If which  factors)  to  can  encompassing  that  some m a j o r p r o c e s s e s For  eliminated,  production?  by  and  systems  systems  i s not  production  due  rate.  is  scale  s i m p l i f y the  phytoplankton  extinction  scale),  to  is  aquatic  purpose here  estuaries.  estuary  effect  other  mechanisms of  such c o m p l i c a t e d  microcosm  The  contrary,  operational  of  a  system.  are  experiments  features certain functions  the  mixing  of  was  the  to  biota,  characteristics of  estuarine  of  two  very  10 2.  Experimental  design  2.1. P r e c o n s i d e r a t i o n The flow  e x p e r i m e n t a l model s i m u l a t e s t h e  estuary  effect).  It  estuary  and  diffusion from  a  entrainment  i s assumed t h a t sea  from  takes place e n t i r e l y  being  negligible,  and  deep  freshwater layer  layer  the bottom  (average  tidal  by a d v e c t i o n .  the r a t e of flow can  influx  remains  i n a two  t h e e x c h a n g e o f w a t e r between  k n o w l e d g e o f t h e mean s a l i n i t y  layers the  with  process  of the  (Pritchard,  almost  Horizontal  be  calculated  i n f l o w and  1 9 6 9 ) . The  unchanged,  the  outflow  salinity  of  only  the  so t h a t  upper  layer  i s considered. It i s  assumed t h a t  the depth  upper  layer  is  steady  of the e s t u a r i n e  constant  circulation.  I f the  considered  as  estuary  at  salinity that flows  out  happening and  from in  these  evolution will  body  rate,  processes  can  estuarine  flows  the  t o t h e mouth o f  salinity  requirement  element.  Thus t h e r e a r e two  body o f w a t e r ; s a l i n i t y  water  becomes d i l u t e d .  also  happen  t h e one  the reflect  system  pattern  of  the s p a t i a l • because  high  major  water  processes time  Since only advection i s  i s i n dynamic over  spatial  mixed  i n c r e a s e s over  space.  the c o n s t i t u e n t s c o n t a i n e d  to  of  the  of the water w i t h i n  of c o n t i n u i t y ,  equilibrium, The  i n the  dimension.  water with entrainment  dilution  an  the  that  a l l  of  state  (see F i g . 1 ) , the entrainment  increases  processes of  which  t h e whole c i r c u l a t i o n  correspond a  speed  that  and  two  element  As  the o r i g i n a l  considered  in  average  element.  the  the  body o f w a t e r a t t h e head o f t h e e s t u a r y i s  one  seawater  at  of  temporal w a t e r mass  T h i s means  o f sea water a t a  temporal  evolution  that  certain of  p a t t e r n of those processes  a l l  the  parameters  carried  the in by  River  S e a  •i i Figure  1.  Flow diagram f o r the upper l a y e r box o f number n i n a two d i m e n s i o n a l model o f e s t u a r i n e circulation with v e r t i c a l mixing (advection o n l y ) . S, s a l i n i t y ; Q, volume o f f l u x r a t e ; Sv s a l i n i t y at l o w e r l a y e r , Qv, volume of f l u x from the bottom l a y e r , below: typical salinity profiles; 0, l o w s a l i n i t y e n d a n d S , high salinity end..  12 this  moving  relative  body  to  p o s s i b i l i t y to  a  of  constant  of  One  point  experiments  is  in  the  laboratory  conditions,  production  system  the  l i g h t  process  and  begins. the  the  But  estuary  beginning  into  us  real  The  estuary  from  marine  phytoplankton the  estuary  ecosystem  system.  The  conditions  light  the  is  not  an  sea  water  deeper  not  start  zone a  sea  of  the  entirely  in in  in  not  for  laboratory.  In  whole  by  the  that to  coincide  features  the  head  of  is  the  drives  the  start)  which  c i r c u l a t i o n  layer.  are  the  suggests  layer,  this  our  until i f  phytoplankton  with  these  growth  entrainment  under  pattern carried  This  photosynthetic  components  of  photosynthetic  biological  underneath  through  primary  operation  phytoplankton  water  water  laboratory point  the  sure  reflect  estuaries.  starting  of  temporal  should  the  the  the  reproduce  The  the  processes  of  to  process.  estuarine  l i m i t a t i o n  euphotic  biological  development  species  the  can  evolution w i l l  of  seawater  because  reach  consider  are  the  desired  activates  we  with  of  equal  of  rest  laboratory  point  evolution  of  us  in  controlled  phytoplankton  that  they  of  an and  energy  analysis  the  of  starting  estuary,  we  major  that  the  a  be  estuary  temporal  (where  of  ecosystem. t e l l s  in  must  Light  state  the  essential  a r t i f i c i a l l y  source.  water  components  is  the  is  of  dilution  these  there  a  e c o l o g i c a l component  which that  ecosystems  of  in  provides  evolution  the  some  distribution  body  entrainment  and  stay  This  settled  temporal  pattern  spatial  both  a  gradient  evolution the  the  actually  carrier.  using  through  s a l i n i t y  water  their  investigate  features  of  the  dominated laboratory  estuary  system.  13 On  t h e o t h e r hand, what  components the  upon  salinity  phytoplankton estuary  may  response  reaching  Does  components  or  actually  the  of  the estuary?  gradient?  be  components.  i s the s t a t e of the freshwater  first  the  this  salinity  other a  living  grave  experiment  freshwater  What happens  for  is  t o them a t  gradient f i l t e r  out  components?  The  some  of  designed  ecosystem  ecosystem  freshwater  to look at the  when i t e n c o u n t e r s  the  s a l i n i t y gradient. A  natural  been  used  known  to  as  (Grice  water analyse  Reeve,  important  tool  of  in  for  reduced  to  ecological  level of  and at  components;  water  chemostats  d e s c r i b e d by o t h e r s  understanding et  studies,  is  the  size  However,  large t o be  for  these  manipulated.  of the container i s  which  conditions  of food chain  volume o f w a t e r i n  and d i f f i c u l t  "microcosm"  as an  a l . , 1978). For the  unnecessarily  the  this i s  has been s u g g e s t e d  (Parsons  study,  a  will.  ecology many  of  the  can  can  be more  more  "batch c u l t u r e "  easily  accurately  be  type microcosms  bodies which can not s i m u l a t e the e s t u a r i n e and t h e e c o l o g i c a l  chemostats  in  our  expensive  conditions  microbial used  too  that  isolated  physical  furthering  enclosure  and this  controlled  hand,  i n a l a r g e c o n t a i n e r has  T h i s method  microphytoplankton  manipulated  are  of  mechanisms  "Mesocosm"  organisms, So,  some  1982).  transformation  an  captured  a "mesocosm" and t h e y have been  &  purpose  column  have  long  (Jannasch  studies  connected  of in  processes.  been u s e d et  as  a l . 1974)  phytoplankton sequence  has  On t h e o t h e r  powerful and l a t e l y  ecology.  been u s e d  tools  in  have been  A series of by C o o p e r and  14 Copeland  (1973)  f o r studying  spatial  salinity  flasks.  I n an e s t u a r y ,  gradient,  i s  continuous  a  of  dampened  an  salinity  2.2  i s  estuarine Our  to i s  the  estuarine  towards  equilibrium,  be b r i e f l y The  dilution  salinity  rate,  of  both  properties  and  the  like  I n terms o f i n a state of gradient  i s  laboratory  ecosystem i n experimental  are only  valid i n  admixture o f sea and f r e s h  systems  a  so t h e nature of t h e  and i n t e r p o l a t i o n s  processes  chemostat  gravitational  steady state.  i t may  field  i n  (e.g.  common f o r t h e m i c r o b i a l  results  common  of  However, f o recosystems, t h e  oscillation.  gradient  (Leffler,  water  1980) and n o t t o  (e.g. sedimentation, e t c . ) .  . Design of the Experiment Since  import  the and  gradient,  can layer  be  experimental  export  the  incorporated  and  ecosystem,  a constant  conditions.  other  gradient  reaching  with  the  which  before  harmonic  associated  tending  of phytoplankton i n a  i n the series  the physical  process  culture  stability  respect  created  etc.) i s i n equilibrium.  estuary  both  gradient  the growth  exchange  into  which  export  continuous  was d e s i g n e d t o s i m u l a t e  characteristics  "openness"  of  the  system  the "batch culture".  considered i n  apparatus  to  be an  import  o f mixed system  more  closely  this  consideration,  of  water  would  (Cooper  water  first  a  system  of a very different  estuarine  between  be  ecosystem water  character  importance.  t o resemble  an approach  to  f o r t h e upper  a l . , 1973; M a r g a l e f , 1967).  adopted  salinity  had  An e s t u a r i n e  a r e o f paramount appear  et I  "open"  and  An open dynamics Based  batch  on  cultur  15 time  around  constant of  days)  input  mixed  over  salinity  containers  volume  of  and  or  was  used  filtered  simulate Every each  over  studying  effects  as a r e s e r v o i r  one,  I  tested  the fresh  the  system  was  sea water  salinity  in  unfiltered the  fresh  initial  Unit  three  flasks  the  water f o r both  held  conditions  two, using  procedure  as  sea  original  that  in  intensity A 20  litre  unfiltered  water  plastic  bags t o  each.  ecosystem with  water  time.  a  was  water  test  followed  and  so that t h e unit  sea water  "control  as t h e o r i g i n a l  water  In  a r e t h e same.  first  In unit  two,  which  decreased over  and f r e s h  one  fresh  the flask  filtered  produce  In the  components.  unfiltered  over  I and I I  stage  light  black  the salinity  to  sea water  be managed by one  o f two f l a s k s  run into  stage  and  i n two s t a g e s . W i t h  water  that  original  environmental stage  so  suitable  i n estuaries.  increased was  size  microflagellates  fluctuation.  was a l l o w e d t o r u n i n t o  water  or  was p e r f o r m e d  initiated  flask  by a  The  represented a  These were covered w i t h  of experiments  filtered  i n the flasks.  constant  to hold  lflasks  an i n c r e a s i n g  could  natural  layer  6  and a c o n s t a n t o u t p u t  bacteria,  and  of  stage having three units  stage,  time  analysis  lower water  set  water  temperature  water.  dark  or fresh  This established  the  the  four,  f o r our experiments for  Constant  bottle  in  gradient  water  eliminated  achieved  time.  used  phytoplankton person.  was  of sea water  water  decreasing of  6  was  time.  Each o f t h e  under c o n s t a n t culture".  water, to  In  t h e same  observe the  i  seawater  ecosystem  b e h a v i o r on t h e s a l i n i t y  gradient  (see Table  1). New  medium  was  pumped  in  through  Tygon  tubing  of 8  mm  16 diameter  at  residence pumped  a  the  One new  collected  litre  day  o f m i x e d c u l t u r e medium  medium  for  r a t e o f 0.I6  (dilution  was  analysis  pumped  and  i n . The  subcultured  - 1  ,  was  outflow a t 24  h  ( F i g . 2)  Because  the  scope  experiments  , many  important were  1 1/day  days).  before  was  intervals.  and  6  time  out  water  r a t e of  and  p a r a m e t e r s were n o t m o n i t o r e d .  essential  measured  were n e c e s s a r i l y l i m i t e d  as  in  size  Only the  most  v a r i a b l e s f o r each ecosystem  indicators  of  the  dynamics  of  component the  growth  pattern. For size and  of  were  and  the  average s i z e  autotrophic  this  cultured there  increase exponential acclimated my  control _in  vivo  method  was  (Kiefer,  "batch  in  cell  a constant the  the  level,  the  biomass.  The  measured  with  1973).  general  The  reason  subsamples needed  cell  in  this  with  accurately  abundance a f t e r  environment  did  average  to  c u l t u r e " c o n d i t i o n s . Even  phytoplankton  fluorescence  community c o m p o s i t i o n and  components  i s t h a t a l l the  of  the  concentration  exponential  reflects  the  c u l t u r e s have  et al.,1981).  Since  c o n d i t i o n s were a l w a y s t h e  same,  not  correlation  (Brand  the  method  environmental  e x p e r i m e n t s have shown t h a t t h e  increase in  number and  s u b s t a n t i a l v a r i a t i o n s per  experiments  experiments  number and  phagotrophic  relative  under  assemblages  of  the  level  method  further are  natural  conditions,  the  f o r non-pigmented m i c r o f l a g e l l a t e s  to o b t a i n the  the  choosing  though  the  At  in vivo autofluorescence  be  in  conversion.  the  of  bacteria, only  b a c t e r i a were m e a s u r e d f o r b i o v o l u m e  determined  biomass  for  the  biocarbon  number  for  heterotrophic  show much d i f f e r e n c e between between  in v i v o  fluorescence  part 1 experimental flasks  control flask  part 2 experimental flasks  control flask  col taction flask  collection flask raservlors  Figure  2. D e s i g n o f t h e e x p e r i m e n t s w i t h  three  units,  EXPERIMENT  Part I storage supply  Part I I  Control  filtered „ sea water (29 °/00)  f r e s h water (0 °/oo)  original water  f r e s h water (0 °/oo)  filtered, sea water C29 °/oo)  control vessels  filtered f r e s h water (0  original water  Stage I experimental  storage supply  (29  °/oo)  /oo)  Stage I I •xperlaental  T a b l e 1. 6tages.  filtered sea water (29 f r e s h water (0 °/oo)  /oo)  control vessels  The p r o c e d u r e s o f t h e e x p e r i m e n t i n t h e  two  18 and  extracted  chlorophyll  inorganic  components,  measured:  the  dilution NH4  +  is  a  Condition  of  and  water  filter  dissolved  substances  the  of  order  bank  fresh  where  the  water  was  divided  one  High 200  part  away  Department covered  with  kinds  was of  reservoir  urn)  a t New  phytoplankton  and  by  into  two  dilution.  was  Fisheries black  filtered water in  gradient.  After  Oceans  through  were s t o r e d  and (15°C)  um  AA  0.8  um  controlled  Millipore  flasks  as  filter. 25 m  laboratory  initial  and  of the  T h i s water  Some o f t h e s e a filters.  i n a p r e c l e a n e d 20  The  unfiltered  immediately t r a n s p o r t e d  Millipore  km  collection,  stored  ( F i g . 3).  .  the  for adaptation.  a t t h e West V a n c o u v e r and  from  a p p r o x i m a t e l y 32  p a r t was 0.8  retained  collected  performed One  and  of the water i n  pumped f r o m a d e p t h o f a b o u t  room  1  populations  i n a temperature  through  plastic  6  was  0.8  particles  i n the s a l i n i t y  days  parts.  with a  pass through. T h i s  a t low t i d e .  was  filtered  fine  Westminster,  f o r two  controlled  or  were  of  by  some  salinity)  experiment  water  of  For  the concentration  chemical properties  (0°/oo  f r o m a dock  temperature also  let  i t s properties  River  was  salinity m  to  (<0.45  stored  room  and  and  o r f r e s h w a t e r were f i l t e r e d  but  water  was  0.98).  i s a process of  both are a l s o a f f e c t e d  from S t e v e s t o n Harbour,  water  concentration  utilization  original  of the F r a s e r  upstream the  the  to investigate  The  =  t o remove t h e p h y t o p l a n k t o n s e e d  microflagellates  much  2  of nitrogenous n u t r i e n t s  NC>3~  of  (r  Experiment  sea  Millipore  linear  utilization  by b a c t e r i a ;  Natural urn  of  result  was  kinds  autotrophic  mineralization 2.3  two  dynamics  and  a  1 Pyrex  was to a  water  These flask  two as  e x p e r i m e n t a l water  19 under also  light  stored  were  The  front  of  performed  experimental  banks  of  of  the  position.  flask  were  and  200-250  smaller  closed  by A  magnetic  a  12:12  LD  (2  ug*at  1~1)  added  thought the  to the  to  be  seasonal  major  ,  water.  ratios  of  f o r the growth  atoms),  to  nutrient marine  to  the  make  for  Redfield  sure  that  autotrophic  environments  ratio  to  (Antia  et  is  al.,  flasks  eliminate were  ug-at  l  enrichment  First,  Second,  -  1  )  was  to decrease  concentrations to  and  create  which  control  incubation  C  the  1963;  the  experiment.  ( e . g . 106  as  The  to  (50  n i t r o g e n would  growth  f o r diatom  of diatoms  t o a p p r o x i m a t e l y t h e same t i m e f o r e a c h  according  of  1  nutrient  reasons.  Third,  middle  (20 u g - a t I " ) ,  silicate-Si  both n u t r i e n t  silicate.  of  A l l experiments  Nitrate-N  Initial  tubes  but p r e v e n t i n g  used  i n the source water.  favourable for  1978).  exchange  sinking.  and  the  irradiance  was  cycle.  variability  nutrient  requirement  air  necessary f o r four  conditions  time  initial  in  (Chan,  stirrer  and  All  controlled  fluorescent  measured  permitting  performed  were  white, _1  was  f l a s k s were p o s i t i o n e d i n  dinoflagellates  space  sea water  a temperature  i s a saturating  over  phosphate-P  liter)  cool  heterogeneity on  15°Cin  uE*m~2s  caps  contamination.  (6  This  began. I n t a c t  covered with black p l a s t i c . at  four,  irradiance  species  the experiment  i n dark r e s e r v o i r s  experiments room.  until  : 16 N be t h e  case Ryther  have  Fourth : 1 P  by  limiting  of  natural  &  Dunstan,  1971). Every duplicates  part under  of  the exactly  experiment  was  run as  t h e same c o n d i t i o n  and  one the  unit  of  two  experiment  Figure  3.  Sampling s i t e s f o r l a b o r a t o r y surface s a l i n i t y d i s t r i b u t i o n plum  e x p e r i m e n t s and in Fraser estuary  and  m °  21 was  repeated  not  to  study  different the  the  same  pattern and  they  qualitative pointed  reaction out  may  exhibit  are  compared  would  that  at  functions,  of  t h e major  variance  existed  times  any  within  two d u p l i c a t e  f l a s k s . Thus  in quantitative  t h e same.  data, the  Haque e t a l . (1980)  i n two i d e n t i c a l m i c r o c o s m s  the analysis  When d a t a  of the variance  functions  eliminate  the  or  power  the  two  spectrum  e f f e c t s o f p h a s i n g . In  same v a r i a b l e s  quantitative i n duplicate  o r i n two e x p e r i m e n t s a t d i f f e r e n t  but the q u a l i t a t i v e f e a t u r e  o f how t h e g r o w t h  and how much t h e d i f f e r e n t p a r a m e t e r s may one  instance  due  vary  t o i n t e r a c t i o n among them  one s y s t e m . Identical  during  of  conditions, may  relatively  b u t be s h i f t e d i n p h a s e .  one i n s t a n c e ,  at  and  t h e two m i c r o c o s m s d i d n o t r e p l i c a t e .  between  dynamics  quantitatively  may  flasks  i n t h e same e x p e r i m e n t a l  i n t e r e s t s h e r e were n o t how much  of the year,  pattern  that  however,  at  form,  autocorrelation  summary,  flasks  be  a n y one i n s t a n c e ,  infer  Calculation  the  some d i f f e r e n c e should  duplicate  compare t h e s i m i l a r i t y o f  some p a r a m e t e r s  similar  likely  between  show  The p u r p o s e was  but t r y t o ensure that  toqualitatively  pattern  may  between  of events occurred  try  evolution  while  variability  seasons o f the year,  conditions the  a t d i f f e r e n t seasons o f the year.  1984-1985. any  dominate  Seasonal  experiments  change the  variations  differences  in  species  a t d i f f e r e n t times  Because o f t h e c o m p l e x i t y o f in  dynamic in  was c a r r i e d o u t  an  biologically state  of  estuary  composition  important  constituents  estuarine  ecosystems.  results as  estuarine  well  in as  dramatic quality  and  22 quantity  of  dy n a m i c  biomass  salinity  within  the  different  gradient  seasonal seasons  assemblages to  the  which  original  conditions?  collected Table at  at  variation?  may  or  The  different  not  o f t h e water  2. S a l i n i t y o f i n i t i a l  used  f o r the laboratory  waters  their  original  water  f o r the experiments  5, 1985 29, 1985 26, 1985 3, 1985 23, 1985  collected  at different  experiments.  o/oo West  0.000 0.000 0.000 0.000 0.000  Vancouver 29.488 28.614 29.958 27.490 26.630  3. Methods o f A n a l y s i s  of  samples  Nutrients As  major  limiting  were  measured.  were  immediately  24  from  o f t h e y e a r a r e a l m o s t t h e same.  New W e s t m i n s t e r  3.1  the  being  controlled  different  of  at  species  pattern  (conditional  be q u i t e  Salinity  Feb. Feb. April July July  seasonal  t h e same g r o w t h  salinities times  collected  times of the year.  Table times  different  Does t h e  on t h e e c o s y s t e m  water  condition  may  2 shows t h e s a l i n i t y  different  less  same  or  interaction.  Does  contain  more  chamber)  level  have t h e same e f f e c t  which  show  subjected  and t r o p h i c  h)  glass  nutrients,  A l l samples filtered  fiber  filters  both  collected through  nitrate  and  i n laboratory  precombusted  ammonium  experiments  ( a t 450°C f o r  (Whatman 934-AH, 2.4 cm d i a m e t e r ) ,  23 stored  in  Nalgene  were  quickly  for  nitrate  al.  (1967)  nitrate (0-4.5  thawed b e f o r e a n a l y s i s .  plus  1984).  3.2  Total  into  filtered  treatment. and  in The  method  (1984).  filtered  solution  of  (Parsons  from  each experimental  numbers.  immediately (37%) .  quickly  and  the  were  distilled 4  incubated  in  fixed  with  thawed  2.5 ml o f  before  formation  further  to detritus  of  bacterial  f o r counting  a  in  M  0.44 M s o d i u m c h l o r i d e sonification one  ml  water  which  had  from  Velje  tetrasodium solution for  was t a k e n  dilution  sample were f i l t e r e d filters  ( pore  size  (in  from t h e  p i p e t t e and p u t i n t o  to get a f i n a l  membrane  purposes.  a t 100'W f o r 1 min  aliquot  an a u t o m a t i c  ml o f d i l u t e d  polycarbonate mm  flask  Samples were s t o r e d i n  a random d i s t r i b u t i o n  Then  25  et  S u b s a m p l e s were  the attachment o f b a c t e r i a  t r e a t e d with  Finally,  Ammonium  s a m p l e s o f t h e b a c t e r i a must be p r e t r e a t e d  sample w a t e r w i t h  diameter  and  and  suspended  experiment).  Nuclepore  1~1.  0.001  my  times.  ug-at-N  Samples  min and t h e n  of  The r a n g e o f  was t a k e n  15  treated  II.  concurrently  bacterial  frustules  have  determination  f o r disaggregation of bacteria  pyrophosphate  and  -20°C  t h e water to  were t a k e n  vials  phytoplankton  order  measured  total  Because  aggregates,  0-50  um) f o r m a l d e h y d e  at  samples  numbers, b i o v o l u m e s and b i o m a s s  of  glass  (0.22  dark  was  o f 10 m l  enumeration  Automated  Autoanalyzer  were  bacterial  Stored  was done a s d e s c r i b e d by A r m s t r o n g e t  Technicon  1~1)  Aliquots  the  nitrite  with  ug*at-N  placed  and f r o z e n t o -20°C.  concentrations  al.,  for  bottles  9 ml  of  10  through  o f 0.2 um  been p r e s t a i n e d 24 h i n 0.2%  o f I r g a l a n b l a c k BGL i n 2% a c e t i c  acid);  afterwards  0.4  24 ml  acridine  and  then  was  carried  an  orange  added t o s t a i n t h e b a c t e r i a  t h e sample was  Depending  filtered.  out with a Zeiss  epifluorescent  9-14  was  on  fields  the  dispersive  and  a  total  counted  (Venrick,  according  t o the equation:  , . Cells/ml •  of  microscope  system state at  1978).  stained  S l i d e s were made and  compound  illumination  f o r 3-5  (Hobbie  Total  equipped  on t h e  with  filter,  200-400 b a c t e r i a  numbers  area of f i l t e r  counting  e t a l . , 1977).  of b a c t e r i a least  min  were  were  calculated  x X x 10 x  1.37  area of counting g r i d x 4 X = mean number o f c e l l s 4 = stained 10  water  = dilution  1.37  =  stained  on t h e f i l t e r  width each  (W)  biovolume  of  the  s a m p l e and  = 2.0  cell  of  (Bratbak,  1985).  literature.  2  by  specific  carbon  (Bowden,  1977;  was  x  Robinson  et  content Watson,  suggested  1 3  of 1979;  was  calculated  (L)  and  o f 50 c e l l s as  in  :  (pi/4)W (L-W/3) both rods carbon g  al.  by B r a t b a k  the length  2  = to  10"  um ,  t a k e n as t h e a v e r a g e  Bacteria  2.0  reported  um~3  applies  um ,  bacteria,  t h e volume o f c e l l s  formula  tetrasodium  2  x 10&  the  was  Biovolume This  and  solution,  area of f i l t e r  the  membrane,  f o r added f o r m a l d e h y d e  a r e a o f c o u n t i n g g r i d = 10,000 For  fields,  index,  correction  pyrophosphate  o f 9 o r more  c o n t e n t was  c  cell  (1982)  the  cell  -  and  1  and  cocci  taken  for  from  the was  biovolume 1  x lO" ^ g  1979);  (L=W)  bacteria  average  i s 2.4  Hagstrom, (1985).  (L>W)  5.6xl0"  C 1 3  to um~3  g  C  25 B i o v o l u m e / ml Bacteria 3.3  The  =  bacteria  carbon/ml = biovolume/ml x carbon  Measurement  of  gradient  nutrient  and  initial  laboratory  collection  of  salinities  for  conservation equation  salinity  were i n our  water.  with  a  measured department the  day  were  continuity  dealing  conversion  calculation  During  each  and  and  volume  of  salinity  d i l u t i o n change  salinities  oceanography  number/ml x a v e r a g e  of  two  in  the  immediately  experiment,  the  physical after  intermediate  calculated  according  salt  the  dimensional  using box  the  model  to  following (Pritchard,  1969): Vl Si  ' Si-! 4  S  0  •  v  2  = volume o f  experimental  water,  V  = volume o f  water  from r e s e r v o i r  2  inflow  Si  =  salinity  of  that  S  =  salinity  of  reservoir,  Q  =  salinity  c a l c u l a t i o n was The  effect  of  by  by  mixing  with  concentrations.  assumed  that  nutrient  identification  of  For  computer  and have  litre),  water,  biota  and  different the  as  program.  concentration  pure  concentration  calculated  Microflagellates methods  two  (1  day,  in nutrient  phytoplankton  t h u s change was  in experimental  done w i t h a s m a l l  nutrient  Many  day  previous  dynamic changes  utilization  3.4  •  Vi  Si_i  and  2  •  Vl  The  V  «  result  bacteria waters  and of  for  dilution different  dilution effect i t  i s a conservative  from  was  property  salinity.  phytoplankton been  reported  microflagellates  (  for  Fenchel,  enumeration 1975;  and  Sorokin,  26 1979;  S h e r r e t a l . , 1983 and Hewes, 1 9 8 3 ) .  was  direct  counting  40xmagnification. drops H2O  of +  were 2  150  Lugol's  190 done  ml  ml  were  because those  the  but  important For  that  of  identified  were  counted  as  one g r o u p .  based  on  trophic  regarded 3.5  In of  vivo  biomass  Samples  time.  of  were  inserted  genus  as b e i n g o f major  Standing  stock of  held  at  identification (volume =  andautotrophic  were  in  the  not  are larger  included  All  in  the  size  between 2-10  than  t h i s may  Ciliates  this  group.  have a  were  suggests  or  Ciliates they  um  found to  were n o t  must  play  a  numbers.  o n l y major dominant s p e c i e s  s p e c i e s , a l l o t h e r minor s p e c i e s  Because I c o n s i d e r e d  level,  the microcosm  the species composition  was n o t  importance.  phytoplankton  i n t o a Turner  and  i n 2000 ml  chamber  heterotrophic  was m e a s u r e d t o g i v e an  phytoplankton  The s u b s a m p l e s  salinity  base p l a t e  flagellate  fluorescence the  and  2  number made i t v e r y d i f f i c u l t  presence  were  system  100 g I  Counting  of phytoplankton  to  +  ( F e n c h e l , 1982b).  in controlling  analysis  KI  microzooflagellate  counts  their  role  of  b u t t h e low  meaningful  counted,  the  function  samples  obtain  g  flagellates  only  in  (200  CH3COOH).  counting,  trophic  at  a c c o r d i n g t o t h e p i n k and brown c o l o u r .  non-pigmented  counted  microscopy  s a m p l e s were p r e s e r v e d w i t h 5  Identification was  light  here  water  solution  glacial  microflagellates  higher  routine  i n a i n v e r t e d microscope  ml).  the  using  The method u s e d  25  x  150  10-000  were t h e n same  assemblages mm  cultured  condition  i n the c u l t u r e s .  culture  fluorometer  indication  tube which  were  e v e r y day a t a g i v e n  i n sequence a t c o n s t a n t and  time  period  as i n  27 experimental and  _in v i v o  then The  flasks.  the  For  the  fluorescence correlation  regression  fluorescence  were  experiment, measured  and r e g r e s s i o n  equation  with  first  for  extracted  in  chlorophyll a  simultaneouslyand  between them c a l c u l a t e d .  vivo  chlorophyll  fluorescence  and t h e  a  acetone  in  90 °/o  was 2  Y = 0.70X + 1.73, ( r = 0.98, N = 6) The  regression  fluorescence  equation  (of extracted  to  phytoplankton for  these can  Bacillariophyceae  Parsons,  a  (ug/1)  and  2  ( r = 1.00, N = 10)  two e q u a t i o n s be  chlorophyll  v a l u e ) was :  Y = 0.167X + 1.74, according  for  calculated are  not  c h l o r o p h y l l a and b i o m a s s o f i n t h e e x p e r i m e n t . The  very  1969; P a r s o n s e t a l . , 1 9 8 4 ) .  different.  ratio  ( K i e f e r , 1973;  28 4. 4.1  Experiment This  stage  question  of  components of  spatial  of  on  I  of  the  what  temporal  water  stage  is  the  the  population  water  was  the  phytoplankton. by  a 0.8  population  and  as  dilution  either  an  increasing salinity  these 4.11  in  part  salinity Increasing  water or  2.  The  gradients salinity  by  water.  salinity  shown  The in  clearly of shows  water  4.  constructed  phytoplankton the  salinity the  Fig.  adding  dilution. algal  as  A small  growth; the  a t the  head  in this  experiment  t o remove t h e  experimental  sea  then  sea sea  water  I used i t  water t o  1 or a decreasing  create  salinity  f r e s h w a t e r components  on  gradient an  increasing salinity  filtered  seawater  during  the  Stage  I, p a r t  nutrients between the  combined  i n c r e a s e on  (29°/oo)  salinity  1.  The  two  effects day  1 and  m a j o r s p e c i e s was  to  NH 4>  period  gradient  of  the  2 was  on  to  was  5a) the  salinity due  is  curve  (Fig.  components  was  fresh  resultant +  (N0~3,  these  gradient  experimental  i n c r e a s i n g temporal  and  gradient  fresh  were i n v e s t i g a t e d .  i n the  interaction  estuary  i s a c t u a l l y that of a  response of  change An  functioning  filter  in part  to  p r e c o n d i t i o n of the  So  initial  In e x p e r i m e n t p a r t one, maintained  um  extrapolation  ecosystem  m i c r o f l a g e l l a t e s and  either  gradient  the  the  f r e s h water e c o l o g i c a l  estuarine  b a s e d on  t o answer  Because the  e v o l u t i o n curve  of  filtered  phytoplankton  gradient. an  designed  of  components w h i c h s t o p  estuary,  water  of is  was  behavior  salinity  distribution  the  experiment  evolution  autotrophic  Results  and  fresh  T h a l a s s i o s i r a spp..  On  Figure  4.  The t e m p o r a l s a l i n i t y p a t t e r n s t a g e I o r I I , p a r t 1 w i t h an gradient.  i n a l l of the experiment increasing salinity  Figure  5a. The d e v e l o p m e n t p a t t e r n o f a u t o t r o p h i c component and n u t r i e n t s i n t h e e x p e r i m e n t s t a g e I , p a r t 1 d u r i n g J u l y , 1985.  O  *-  2.5  .-A  o 2.0 1  1.5 E 3  C w  E E  <  1.0 0.5 0.0  I  T"  0  5  Figure  5b.  The d e v e l o p m e n t experiment stage  T  10 D a y s  15  +  p a t t e r n o f NH 4 i n t h e I, p a r t 1 d u r i n g J u l y , 1985.  20  0.30 4 0  1-0.25 g  Z 30-1  -  1-0.20 J hO.I5  20  | u.  I-0.10  ? 104  f-0.05 «5 10  15  20  Days Figure  6 a . The d e v e l o p m e n t p a t t e r n o f a u t o t r o p h i c component and n u t r i e n t s i n t h e e x p e r i m e n t s t a g e I , p a r t 1 d u r i n g F e b r u a r y , 1985.  0.00 K  o 1 3.  E  2.0 1.5 1.0  %  4  O  E 0.5 <E 0.0  T"  T  5  15  10  —i 20  Days Figure  6b.  The d e v e l o p m e n t p a t t e r n e x p e r i m e n t s t a g e 1, p a r t  +  o f NH 4 i n t h e 1 during February,  1985.  34 day of  3,  when  algae  diatoms  were died  (Blanc  changes  (Qasim, vivo At were  the  1972;  group  of  10-20°/oo  (Blanc 1975).  also  the  10°/oo  showed  very  i n c h l o r o p l a s t s ) and  auxospores  decreased  with  The  6 was  caused  An  diatom  was  It  may  have been  (unidentified).  The  15  have been due range  t o 25 °/oo  freshwater  was  contaminated  experiment  Mahoney, carried  1979;  out  decrease of n u t r i e n t s rapidly of  during  the  N0~3 out  t a k e n up  and i t  influenced  by s p o r e s o f around  Qasim,  i n February  1972; showed  (Fig. in  5b,  the  6b).  Even  F e b u r a r y , 1985  concentration first  period  time  ( F i g . 6 a ) . Ammonium w h i c h  than n i t r a t e  of  decreased though  t h e r e was +  NH 4  to  i n which  have an o p t i m a l g r o w t h s a l i n i t y 1969;  in in  and  salinity  original  increase  spp. o c c u r r e d  (>25°/oo). grew  salinity  by a m i x e d p o p u l a t i o n .  Thalassiosira  The  physiological  increasing  16 may  al.,  water  structural  the  et  carried  phytoplankton, decline  some  p o p u l a t i o n a t day  which  reached  diatoms  1971).  diatoms  growth  other  that  The  more  decrease  experiment  day  the  o f p h y t o p l a n k t o n d u r i n g the whole e x p e r i m e n t a l  no  frequently  and  by a s m a l l r o u n d  saltwater.  increase  the  (Foester,  growth,  salinity  species  and  after of  estuarine  no  h  salinity  Paasche,1975; W e t h e r e l l , 1961).  therefore  Paasche,  of  15°/oo,  Most o f t h e f r e s h  when  changes  decrease  of the l a t t e r  higher  the  6.  day  Some  48  replaced  second  until  diatoms  beginning  appears by  a  within  these  then  decline even  as  fluorescence the  t o about  depressed  morphological  (such  of  were  increased  al.,1969).  formed  state  depressed or  et  distinctive  were  the s a l i n i t y  is  before in  the  no g r o w t h o f  still  of the experiment  showed  a  ( F i g . 6b).  35  This  indicated  autotrophic In  appeared  at  this the  may  was  range  of  in  the  two  In  the  1957;  Parsons  of  found  of  the  two  identified  to the  the  and  1980).  micronutrients  nutrient  rich  phytoplankton.  However,  14)  water comparing  dynamics,  the  p h y t o p l a n k t o n g r o w t h was  dilution  (i.e.  essentially  water  in  an  halophobic  changes release  occurring of  heterotrophic of  (1978)  this  discussed  is  in  on next  the  low  material would the  surface  Dodimead,  rich  in  Sunda, 1 9 7 7 ) . the  B12 The  growth  of  depressed stress.  the  and  nutrient  delayed  Therefore,  species.  that  mechanism o f  oxygen  one  as  the  freshwater  t o a mass m o r t a l i t y o f  which  r e s u l t s from the  salinity and  functions  in  filter  cause  region;  its  the  fresh  osmotic  concommitant  assimilation  decreased  heterotrophic  section.  of  a  due  phytoplankton  microbes  i n the  a l l the  species  Strait  and  considering  gradient)  suggested  estuary  organic  process  out  and  osmotic  salinity  filtering  Morris depletion  the  by  but  themiddle  i n the  stimulate  two  f r e s h water c o n t r o l ( F i g .  experimental  flask  level,  in  is  spp.  species.  major  (Tully  the  These  the  nitrogen  and  the  experimental  in  (Cross  with  the  i s the  water  flask  living  species  River  the  free  range.  groups  bloom  should  to  different  two  estuary  to  due  Thalassiosira  salinity  phytoplankton  al.,  due  entirely  River  not  that  from seawater entrainment  et  1973)  I  dominant s p e c i e s  Fraser  i s supplied  was  most l i k e l y  between  This  spring  layer  inflow  to  connection  annual  (Cattell,  ends  t h e y were n o t  no  which  was  work,  salinity.  Georgia.  and  belong  Unfortunately there  consumption  component  bacteria.  groups  a  O2.  component  The  by  the  effect will  be  36  9 20 M  K  8 15  I '° to  aof 1.0  i  ao  o  10  15  Days Figure  20  7. The development p a t t e r n of h e t e r o t r o p h i c b a c t e r i a and n a n o z o o f l a g e l l a t e s i n experiment stage I p a r t 1 d u r i n g J u l y , 1985.  300 250  1 ©  u e 0  o  200 150 100 500 0.00 OO  10  Salinity Figure  8.  Growth o f different  30  20  To  <%•)  f r e s h w a t e r b a c t e r i a on salinities.  p l a t e media  at  37 Heterotrophic Bacteria, ecosystem, material  as  as  forms;  has  a  V  a very  Van-Wambeke,  10  Watson's of  5  4  work  the  They in not  appeared  Bodonidae very  10^  x  4  Haas  to  (Parsons  2-10 um.  provide  which  Webb  1980).  the b a c t e r i a with their  ecological  a r e i n t h e range consistent  with  6.29 x 1 0 ; W a t s o n , 1 9 7 9 ) .  is  Most  6  shaped a t t h e b e g i n n i n g .  consumers o f b a c t e r i a i n p e l a g i c  t o Kudo  (1979), they (Sherr  1982b).  Most o f them  The i d e n t i f i c a t i o n  out but according  and  et a l .  1984; H o l l i b a u g h ,  fall  o f s p e c i e s was (1966),  are mostly  and S h e r r ,  Fenchel  Monadidae,  1982).  They a r e  i n t h e a q u a t i c e n v i r o n m e n t ; u s u a l l y f r o m 0.4 t o 13 in  various parts of the world  1981).  T h e i r d e n s i t y i s c l o s e t o one.  toward  the  (Sorokin,  Active bacteria  i n our experiment  major  Amphimonadidae  cells/ml  distribution  cells.  complement  cells/ml,  10  organic  1969; Haas a n d Webb, 1979; F e n c h e l ,  carried  and  abundant  lO^  the  range  thoroughly and  others  numbers  an  i n n a t u r a l s e a water i n  o v a l s h a p e d and b i f l a g e l l a t e d .  size  (1982b)  x  are  (Lighthart,  the  with  to  in  particulate  substrate  a r e s m a l l - s i z e d and c o c c i  Zooflagellates systems  x  (1.5  cells  Landry,  capability  component  (Ammerman, 1984; H a g s t r o m e t  1985;  combined  to  They e x i s t  high growth r a t e  The b a c t e r i a l  x  and  o r dormant b a c t e r i a l  physiological  4  heterotrophic  f o r uptake o f o r g a n i c  m  Theseproperties  role.  component  compounds  energy sources.  high  al.,1984;  lowest  organic  active  and  the  the  utilize  two  1977)  and p h a g o t r o p h i c  small was  1981).  size more These  fraction even  The s i z e  (2-4  um)  (101.3-198.3  flagellates  oceans  may  distribution i s  and ug  grow  (Sorokin,  wet  the  biomass  weight/ml)  a t 1.5 x 1 0  2  -  38 2.2 &  x  10  2  Sherr,  cells/ml/h 1982).  natural  These  waters,  responding natural  to  the  cells/ml)  and  high  these  towards  1980a,b).  This  be  due  water  and  salinity 8).  sharply.  the  increase or  population  can  is  no  salinity. be  due  to the  stimulating  are  (Sherr  high  for  capable  of  populations  under  The  x  in  release  products  by  the  due  day  by  the  which  is  6  saline  the  on  optimum ( Fig.  declined  possible explanation  is  the z o o f l a g e l l a t e a  zooflagellate  dilution. of  Unfortunately,  z o o f l a g e l l a t e s to  in bacteria population  e x t r a c e l l u l a r organic  p h y t o p l a n k t o n growth  of  may  impact  l a b o r a t o r y work  situation,  tolerance  day  inflow of osmotic  strongly affect  increase  to  population.  period, microzooflagellates  may  5  (Fenchel,  2 and  i t resulted in  c l e a r ; one  10  lowest t h r e s h o l d  bacterial  substrate  i n d i c a t e d i n my  the  of  1 are  (4 x  concentrations  numbers on  s t r o n g l y a f f e c t e d by  second  day  and  clearly  counts  b a c t e r i a from the  limited  on  been  cells/ml)  limited  addition  i s not  a  low  10-*  represent  of organic  salinity  bacteria  have  b a c t e r i a on  (13.8  the  In  work The  7).  in bacterial  reason  be  quite  between  relations  a substrate  initial  in  population  there  on  release  The  of  may  b a c t e r i a as  During  the  point  organisms. for  bacterial  big b a c t e r i a at high  enrichment of the  freshwater  size  increase  to  of  (Fig.  grazing  grazing  slight  are  microflagellates  trophic  data  small  selective  The  times  interaction  in  zoof l a g e l l a t e  flagellate  doubling  9.7-18.2 h  at c e r t a i n rate.  nanozooflagellates in  times of  dynamics  qualitative  reflected  doubling  indicating  conditions The  with  seems  products or  (Bell  and  to  some  Sakshaug,  39 1980).  Because  the  compared  with  consumed  a carbon  that  phytoplankton. autotrophic  of  Even  of  The  component  makes  The  highest  value  The  average  size  very  3  V = l/6piD the  carbon  the  4.5  um  value  f o r the  (14.1  x  ugChla/1; et for  al.,  10  activities flux  i n the  just  carbon  of  has  cells  the  carbon  standing  (Fenchel,  from  of  the  a  high  have  t h e r e a p p e a r s t o be  condition,  heterotrophic  stock.  1982a)  and and  per  314  4.1  flow.  ug C / l .  um.  If I  then 3  um /cell =  0.75  x 10  - 1 1  1982b).  standing  gC/cell  So  highest  106  factor  ugC/1  three  4.32  (Parsons,  stock only  accounts  components'  idea of the  a more m e a n i n g f u l  energy  index  than  the h i g h growth  rate  zooflagellates  day  for  a maximum was  standing  the  the  s t o c k was  conversion  Nevertheless,  1984)  was  ca.  r a t e of a l l the  w o u l d be  population  a  three trophic  the  carbon  been m e a s u r e d . I have no  (Ammerman,  growth may  chlorophyll  phytoplankton  however t h e  not  entirely  among t h e  (Fenchel,  The  system which  bacteria  =36.1  i s u s e d as a C / c h l  ugC/1;  bacteria  conditions the  30  1969),  129.6  3  cells/ml).  if  that  stress  standing  spherical  zooflagellate  3  have come  carbon  coefficient  diameter  t h a t the b a c t e r i a  z o o f l a g e l l a t e s was  = l/6pi(4.1)  conversion  low  l a r g e c o n t r i b u t i o n to the energy  the are  a was  budget.  of b a c t e r i a l  assume t h a t a l l c e l l s  not  osmotic  organic  of  seems  assume  t h a t under t h i s a  i t  organic products,  of  indicates  we  under  carbon  division  levels  w h i c h may  though  excreted  i n terms of the  stock of c h l o r o p h y l l  bacteria,  source  component  proportion gap  standing  under  high grazing rate Fenchel,  1982b;  certain (20%  of  and  a  40 bacterial  populations  of  cells/day  in  10  a  7  Wright,  1983),  greater  potential  flow  in  of  the  x  both  capability  medium,  species  Tintinnidae  was  transformation  the  in  probably  the  sharing  the  the  functional  x  components may the carbon  have  higher  level  in  h i g h abundance  the (main  level  stock of  r a t e and  6  energy  existed  standing  10  estuary;  implies a high  dynamic  more  i n an  as a t h i r d  a relatively This  t o 7.5  of  carbon  physiological  significant  information for  the  s t r u c t u r e and  e n e r g y pathways  in  ecosystem. results  not  play  the  original  the  gradient  e s t u a r i n e e c o s y s t e m as  water.  They  cannot  changing producer,  analysis  was  package, matrixes  experiment  from  7  interrelationships  between  correlation phytoplankton,  of of  but  an  role.  and  by an  calculated  variables  of v a r i a b l e s . variables  some NO~3,  are  variables +  NH 4  and  through  they the  can  did in  salinity  Their ecological  role  is  organic contributor.  facilitated  were  pass  and  between t h e d i f f e r e n t  pairs  pairs  their  "SYSTAT",  Correlation  comparing  t h a t f r e s h water phytoplankton  i n an  as a p r i m a r y  software  suggest  same r o l e  without  Data  by  two  1982)  spp.).  contain  of  Ciliates  (Banse,  Thus  population  that these  reaching  1978).  These  not  and  cells/ml/day  to c o n t r i b u t e to  interaction  (Heinbokel,  10^  cells/ml  suggest  experimental  revelation  6  x  whole ecosystem.  trophic  state  2  The  the  of a  statistical  IBM  personal  for  each  20  computer.  part  of  observations.  v a r i a b l e s were correlation  shown such  use  in as  bacteria  The  assessed  coefficients  Table  3.  standing with  the  The stocks  time  can  high of be  4\  T a b l e 3. C o r r e l a t i o n m a t r i x o b t a i n e d f r o m t h e 7 v a r i a b l e s X 20 o b s e r v a t i o n s i n experiment stage I, p a r t 1 during J u l y , 1985, between t i m e ( d a y ) , F l u o r e s c e n c e ( f l u o ) , N0~3, NH 4, F l a g e l l a t e s ( f i g ) , B a c t e r i a ( b a c ) , and s a l i n i t y . P r o b a b i l i t y : * p<0.1, ** p < 0.05, *** p < 0.02, **** p <0.01. +  1 | DAY FLUO N03 NH4 FLG BACT SAL  Table  DAY  | 1.000 | 0.903**** 1-0.726 j-0.688**** 1-0.244 | 0.857**** | 0.952****  |  N03  I  NH4  I  FLG  BACT  |  | 1.000 | 1 -0.835****1 1.000 ! 1 I -0.608****| 0.626**** 1 .000 0 .285 1.000 | 0.040 | 0.027 0.723****1 -0. 760**** -0 .757**** -0.397 |1 .000 0.740****| -0. 580*** -0 .658**** -0.475*10 .845****  4. C o r r e l a t i o n m a t r i x o b t a i n e d f r o m t h e 7 v a r i a b l e s X 20 o b s e r v a t i o n s i n experiment stage I, p a r t 2 during J u l y , 1985, between t i m e ( d a y ) , F l u o r e s c e n c e ( f l u o ) , N0~3, NH+4, F l a g e l l a t e s ( f i g ) , B a c t e r i a ( b a c ) , and s a l i n i t y . P r o b a b i l i t y : * p<0.1, ** p < 0.05, *** p < 0.02, **** p <0.01.  1 I DAY FLUO N03 NH4 FLG BACT SAL  FLUO  DAY  I 1.000 I 0.917**** 1-0.967**** 1-0.859**** I 0.036 | 0.868**** 1-0.948****  FLUO  N03  NH4  FLG  1  BACT  | 1.000 | -0. 855**** 1.000 | J -0. 909**** 0.785**** 1.000 0.052 -0.051 0.219 1.000 | 0. 930**** -0.835**** -0.837**** -0.026 I 1.000 -0. 782**** 0.967**** 0.774**** 0.078 1-0.765****  42 easily  explained  However,  if  surface  layer  interpreted  we  in  that:  marine  an  (1) t h e  zone,  on  a  nitrite  nitrogenous  at  head  only begin  and  can  of  the  be  s i n c e time  is  This  f u n c t i o n i n g as  e s t u a r y , and  to function after  (2)  i n the  reaching  the  a  were  negatively correlated  because  the  phytoplankton  nutrients to synthesize  n i t r o g e n as a n u t r i e n t  up  limited  to  mainly  organic  source.  experiment,  substrates  standing  bacterial  The  organic  (Rheinheimer,  1977;  Hoppe,  1982).  two  completed  which  such  u p t a k e may  s e r v e as  the  because a t the  by  bacteria  1978;  Larsson  on  several  correspond  interaction  by  growth  and  Hagstrom,  trophic  very c l o s e l y during  The  were  of  organic  the o n l y  process.  cycles  correlated  as  bacteria  even t h o u g h t h e i r  the c o v a r i a t i o n  components  and  be  bacterial  beginning  released during  Microflagellates  correlated,  clear  stock  products  utilized  very  but  nitrogen  Bacteria also  p o p u l a t i o n s were l i m i t e d  are  significantly  source,  remove  biomass w i t h  incorporated into protein.  phytoplankton  Wolter,  new  H e t e r o t r o p h i c b a c t e r i a were p o s i t i v e l y  phytoplankton  substrate.  these  scale  the  ammonium  take  are  spatial  the  being  the  correlations  f r e s h water organisms stop  compounds  with  moving s e a w a r d a t  a r e t h e m a j o r a u t o t r o p h i c component  but  chlorophyll  energy  these  components.  zone.  Nitrate, with  of these  d i s t a n c e d u r i n g t h e movement o f t h e w a t e r .  phytoplankton  euphotic  evolution  estuary,  components  transition  time  occurring  to  autotrophic  the  c o n s i d e r a body o f w a t e r  as  proportional assumes  by  of  source 1979; not  interactions  c y c l e s between  and the  often,  having  experimental  Pigure  9. The temporal s a l i n i t y p a t t e r n i n experiment stage I t> I I , a r t 2.with a d e c r e a s i n g s a l i n i t y g r a d i e n t  Relative  P o o  Fluorescence  g  J  .  s  P to o  fi  o  -J-  J.  ua c n n>  o  a.  o Oi  Q) 3  •-3 3" a>  a  3 C rr  a  CD 03 f0 < •"I 01 3 rr 0 Qi CO TJ 3 m 3 3 3 rr rr  h-  CO  3  0)  Q C  Q  rr  TJ  rr rr ft> >1 3  t—' t—l  •< -  •»  TJ  h-> fl) o *fl T oo rr ui • fo  fti  rr  zr  • ro C  Hrr 3"  rr O  O,TJ  (D O >-» 0)  0) C rr O  3"  a> oo  3 03  3 3 fO 3 rr  TJ  O  o  —-I  .cj o  1  o —  Nitrate  1 —  01  in>  ©  1  ro m  oi o  1  ( p g - a t / l )  1—  °  Days Figure  10b. The d e v e l o p m e n t p a t t e r n o f NH 4 c o n c e n t r a t i o n i n Stage I, p a r t 2 w i t h a d e c r e a s i n g salinity g r a d i e n t i n J u l y 1985. +  o  D a y s Figure  1 1 a . The d e v e l o p m e n t p a t t e r n o f t h e a u t o t r o p h i c c o m p o n e n t and n u t r i e n t s i n S t a g e I , p a r t 2 w i t h a d e c r e a s i n g s a l i n i t y g r a d i e n t i n F e b r u a r y , 1985.  1.2 ^  I  ~  1.0 1 *  0.8 0.6 H  0  * *  *4  £  | o | <  0.4 0.2 T -  0.0  5  10  15  20  Days Figure  l i b . The d e v e l o p m e n t p a t t e r n o f NH 4 c o n c e n t r a t i o n i n S t a g e I, P a r t 2 w i t h a d e c r e a s i n g salinity g r a d i e n t i n F e b r u a r y , 1985. +  —i  48 time, the c o r r e l a t i o n s o f f s e t 4.12 D e c r e a s i n g In  the  filtered and  second  (0.8  fresh  ecological  um)  culture  day).  increased  quickly  factor  stock 30  showed  some be  was  same peak  decreased.  seawater  inhibited  The  inflow  fresh  as a i n o c u l u m  gradient  (Fig. 9).  not  was  of  was  in  d a y 6 ( F i g . 1 0 a , 1 1 a ) . The  phytoplankton  for  about  as t h e  (chlorophyll  a)  1.88 ug C h l a / 1 a t d a y 13-15  terminated.  healthy  Phytoplankton  carbon  56.40 ug C / l ( c o n v e r s i o n cells  under r i c h  nitrogen  I n my e x p e r i m e n t t h e g r o w t h o f  under a  increased  biomass  a maximum o f  t h e impact o f s a l i n i t y  normal c o n d i t i o n so t h i s  change.  conversion  deviation. carried  pattern by  phytoplankton  a s t a r t e d a f t e r d a y 6 (10°/oo on t h a t  was  a large  experiment  fluorscence salinity  is  cells  the  the  salinity  Calculated  reached  may c o n t a i n  i t .  Freshwater  ( A n t i a e t a l . 1963).  condition  The  decreasing  reaching  which  phytoplankton  value  in  growing u n t i l  experiment  conditions  The  contained  rate  decreased.  standing  into  gradient.  growth  the  I, 5 1 of  b a c t e r i a i n t h e sea water a l s o encountered a  salinity  until  pumped  chlorophyll  The  stage  water  salinity  in  experiment,  was  d i d not s t a r t  increase  the  water  gradually  heterotrophic  decreasing  of  s e a w a t e r was u s e d a s t h e e x p e r i m e n t a l  component a  other.  gradient  part  water  encountered The  salinity  each  day  out during  except 2-3,  February,  there  which  was  an  1985. I t jln  then decreased  vivo  as t h e  T h i s may have been c a u s e d  by t h e g r o w t h o f  a u t o t r o p h s w h i c h were l e s s t h a n  0.8 um and c o u l d  by a d e c r e a s i n g  salinity.  Pigure  12. The d e v e l o p m e n t p a t t e r n o f b a c t e r i a and n a n o z o o c l a g e l l a t e s i n Stage I, p a r t 2 w i t h d e c r e a s i n g s a l i n i t y g r a d i e n t i n J u l y 1985.  a  50 Table  4  variables  in  most  of  shows the  the  directly  example,  except related  ecological  are  because  is  during  given  relation related  of  Hobbie  and  bacterial  in  uptake this  this  of  et  is  not  a l . , 1985).  that  closely  The  and  carbon  and  the a l g a l  also In o u r  flora  state may  the  and  the  energy  (Fogg of  have an  experiment,  the  the  trophic directly  the  potential  research  has  phytoplankton  production dissolved  bacterial  The  relationship  Recent  provides  other  relationships.  development of in  the  variables  and  a close  production  results  stimulate  physiological  microflagellates  other  suggests  d e p e n d s upon many f a c t o r s .  composition the  of  to  each  for and  experiment.  with  organic substances.  biomass  distributions pathways  this  primary  relationship.  which  the  correlated and  microecosystem  substances  of  (1977) d e m o n s t r a t e d  of  bacteria  with  of  which  correlation;  relates  p r o d u c t i o n i n terms of t r o p h i c  Rublee  phytoplankton  and  not  time  correlated  requirement  phytoplankton  confirmed  salinity  course  the v a r i a b l e s  and,  8  between  of the development course of  nanozooflagellates  to primary  between  correlated,  were  time  All  closely  and  correlation  o t h e r show a c l o s e  because  between  e x p l a i n e d by t h e d e v e l o p m e n t  phytoplankton,  the  nanozooflagellates  be  The  and  Time  it  p a r t two.  can  t o each  components  phytoplankton.  coefficient  zooflagellates.  nutrients  phytoplankton  correlation  experiment  parameters  phytoplankton, are  the  of  organic  populations.  The  flow  between  these  two  These  i n c l u d e the species  e t a l . , 1965;  Wolter,1982)  population  (Sharp,1977);  effect  ( W o l t e r , 1982;  possibly  a l l of these  Goldman factors  51 affected The  the  production  production  source by  is  relative  products.  primary  the  (45.40  ug  EOC  full  -source  The  interesting  maintained  of  contribution  to  sustain organic thing  the  level  substrate with  may  um  have  been  the  (Gak  a l . , 1972;  inflow  have  of  occurred  slight  the  the  and  initial  the  an  and  (8.3  x  10^  I, p a r t  one.  of  the  w a t e r was  filtered  This  would  Wright, 1984).  With  z o o f l a g e l l a t e s would  have grown v e r y  b a c t e r i a showed  increase  have  nanoflagellates  b a c t e r i a medium.  may  organic  microflagellates  b a c t e r i a by  higher  population  then  utilized.  bacteria  stage  medium.  Azam, 1980;  microflagellates increased, decrease  some  was  1  boundary  inoculum, the  in a relatively  day  bacterivorous  from  water  So  covariance  of  i n the  upper  Fuhrman and  fresh  at  that  e x p l o i t a t i o n of  microzooflagellate the  an  Most o f  very  bacteria.  density  because  grazing  close  is  the  phytoplankton  requirement.  development  eliminated  released et  the  represent  filter.  that  so  not  microflagellates  comparing  limitation  a 0.8  would  bacterial  carbon  a  Bacterial  higher  gradient,  is  shows  was  p h y t o p l a n k t o n can  likely  and  photosynthetic  must have been  Figure  cells/ml)  biomass  substrates  b a c t e r i a and  12  of  standing  that  (EOC).  a b a c t e r i a l food  stock  salinity  not  between p h y t o p l a n k t o n and  microflagellates.  As  the  is  carbon  of d i f f e r e n t  experiment the  It  as  standing  release  Heterotrophic  This  the  spectrum of  C/l).  can  other  new  its utilization  r a t e of  production  released  the  and  However, i n t h i s  through  small  EOC  e x t r a c e l l u l a r organic  d e t e r m i n e d b o t h by  the  pass  of  of  to a small  Thus  the  quickly.  a peak.  During  this  period,  enrichment of  by  fresh  water  the  organic  water  day. day  we  could 5  assume  supply  were  cells/ml  10^  10^  removed  then  the  by  the  sudden  5  on  day  is  Moiseyev  bacteria. removed  If  similar  with  cells/ml. pathway  the  2 under a c e r t a i n  bacteria  cells/ml. 2,  to  suggesting  the  as  cells/ml  This can  (1983) by  be But  very  10^  can  be  in  the  ecosystem  the  8.4  day  into  justifiable,  10^  carbon  significant  its  Kopylov  times of  daily  2.  explained  biomass  others.  x  from  c e l l s / m l on  is  of the  and  B/F.(Laak  increased  doubling  fast  the  my  t h a t o b s e r v e d by  indicates that  If  x  i s i n agreement w i t h  and  0.67  to  production  of  c e l l s / m l were  z o o f l a g e l l a t e s from a p o p u l a t i o n  evidence  10  10  x  7.5  6 x  produce  calculation  observed  100 may  bacterial  50%  that  population.  flagellates  flagellate  population  However o n l y  i s taken  18.5  a consumption of  day  processes.  one  day  of  this  observed  Wright per  biomass d o u b l e d d u r i n g  grazing  i n c r e a s e of n a n o z o o f l a g e l l a t e s  (1980)  days  one  bacterial  to  salinity  during  by  This value  1  the  be  from  number o f  high  biomass  1 t o day  factor  transformation  magnitude  other  x 10  The  microzooflagellates.  this  yield 10  data.  cells/ml  Thus  1.44  at  to  may  then  x 10^  zooflagellates/ml.  experimental x  16  present  were  al.,1984),  f r o m day  due  f r o m mass m o r t a l i t y  i n t o the  biomass  rate,  to  bacteria/zooflagellate et  actively  bacterial  that b a c t e r i a l  increased  cells/ml  flow  the  zooflagellate  substrate have  10^  x  of  (Ammerman e t a l . , 1984)  organic  grown  substances r e s u l t i n g  Some  into  If  have  o r g a n i s m s when t h e y  (29°/oo).  transformed  10  b a c t e r i a may  of  flux  7 x  10^  through  in proportion  because of  the  to  fast  Nitrate _  -  0  o» 1  .  O I  (^g-at/l) N  Ul I  Bacteria  N  O 1  Oi I  I  No. (xlO*) ^ j  Relative  •  Fluorescence  * oi i  o  Bacteria  o I  — 1  Relative o o  (xlO ) 6  ro  Oi  L .  L>  45—I  Fluorescence o — — ro  C  0 3 ^ C >-3 ill T (C 3 a) a> 01 3 CD t— rr < Ml D) H  "1 3 O 0> CLTJ 0)  3  3" O" fD € 01 3 0) O rr rr rr fD (D TJ 1 1 D) m- r* a a» rr  c  0>  h- 3 3 3  iQ 0*  t4 o  con \-> 3 3"  "< n- a> - *1 O 01  M MC vO rr ao it o Ui Qi rr  •  n  CO  O  W  3"  >< TJ rr h(D O 3  o <o  h. 3 rrTj 3- O 3  o  a>  3  rr  ro  o  (x>  Nitrate (/jg-at/l) •  7S  o  55 uptake  rate  and  exhaust  their  maximum  growth  two  long  and  However, also  likely is  products.  conditions, organic  most o f t h e  substances  source  stimulant  b a c t e r i a may  be  DOC  limited  to  population  shows  4-5.  indicates  source  limited  a  and  by  on  (1)  9.4  2 x  3  response to the that t h e n was  this  photosynthetic 314  ug  C/l  and  under  stressed  were r e l e a s e d  may  brought  produce  bacterial i n the  T h i s may  as  some  use  of  absence  of  suggest  that  required nutrient.  reached 10  i t is  bacterial  f r e s h water a l s o  unavailable  some o t h e r  Fig.10a).  into  that  products  phytoplankton.  day  (see  was  r e a c h e s 46.40 ug C / l .  actually stimulate be  is a  increased  biomass,  biomass i s  of  These  of b a c t e r i a  of  (2) p h y t o p l a n k t o n  w h i c h may  r e l e a s e d by  declined  This  inflow  which can of  release  photosynthetic  general,  limitation  bacteria  carbon  explanations:  the  ( growth  increase  biomass o n l y  material, or,  Microflagellates then  the  in  experiments  incorporated  bacterial  and  the  in chlorophyll a  carbon  from  possible  materials  another  the  of phytoplankton two  in  number o f  the  and  can  potential capability  both f a c t o r s the  bacteria  short periods.  their  phytoplankton  maximum  some o r g a n i c  other  of 6,  the  over  results  increase  that  The  are  The  totally  maximum  There  the  timeperiods  occur  coincidentally,  with  b a c t e r i a , the  to explore  day  calculating  biomass  only  effect  From  and,  not  the  will  limited  the  grazing).  simultaneous  over long  period.  of  constantly  the  rates  time  combination  growth r a t e of  substrate  components may  over  in  great  18.5  x 10  cells/ml.  3  cells/ml  This  decreasing  i n c r e a s e of b a c t e r i a at  flagellate  a l s o removed by  population other  and  was  causes.  day food  There  56 are  two  obvious  population large  may  the  be  ciliates  experiment.  causes  f o r the  strongly  such  as  been  (Spittler  thenanozooflagellates  affected  :  ( D a  by d i l u t i o n  Tintinnidae  They may have  flagellates  decline  and (2) some  s p p . were common  be r e s p o n s i b l e  1973; S t o e c k e r ,  increased,  non-growing  i n the  f o r the decline i n  1 9 8 1 ) . on d a y 15, as  the bacteria decreased.  Summary o f e x p e r i m e n t S t a g e I Both in  f r e s h w a t e r a n d s e a w a t e r p h y t o p l a n k t o n grew v e r y  the controlled  2.  There  zero.  was  13,  other  research  the  sea  1 4 ) . The  water  fresh  were  fresh  quite  different;  concentrations  NO"3/l;  sea  systems developed very  similar.  bacteria  in  the  freshwater  exponential that small  of  the f i r s t  growth  the  phase  not  In t h e  24.19  But  of  the  system  part  s p e c i e s was  a i n s e a w a t e r and  (fresh  ugNO~3/l).  water,  However,  and t h e development  developing showed  pattern  of  an  interesting  of the experiment  (e.g. during  the  completely  i s attributed to  different  p h y t o p l a n k t o n ) compared  s e a w a t e r s y s t e m . The b a c t e r i a l  increase,  different.  was m a i n l y due t o t h e  p h y t o p l a n k t o n blooms  were  during  this  being  water,  patterns  difference  species  water, t h e major  The maximum c h l o r o p h y l l  with  Dominant  90% o f t h e b i o m a s s  In  spp.  nutrient ug  almost  limitation  c o m p o s i t i o n was i n a g r e e m e n t  1984; S t o c k n e r , 1 9 7 7 ) .  costatum.  water  initial  species  (Spies,  control,  Thalassiosira  both  no l a g p h a s e ; N0~3 d e c r e a s e d s h a r p l y t o  f r e s h water and s e a water a r e t o t a l l y  Skeletonema  8.83  almost  and e x p o n e n t i a l g r o w t h a p p e a r e d a t d a y  The d e c l i n e o f b i o m a s s was c a u s e d by n u t r i e n t  (Fig.  in  flasks  well  inhibited  population by  the  with  showed a  growth  of  57 autotrophs of  them  This in  as were  may the  two  been due  Not  periods in  both  process.  The  freshwater  species  salinities  higher  fresh  water  salinity  or  water  adaptations However, estuary water  concentrations adaptation  water  and  and that  heterotrophic  Data  but  continue  may  for  may this  may  still  and  in  may  and  dilution  indicate that  start  growing  function  of  or  be  result  on  predated. some  (1961).  Fraser  River  days.  Fresh  salinity  inhibit  species  i n high m o r t a l i t y .  function  to  convert  heterotrophic which  the  cellular  work  t h a n 10  may  that  after  the  at  component;  through a l l the  impact  components  to  cultured  less  go  (except  also  Wethell's  of  sudden  lag period  release  m i x e d e s t u a r i e s s u c h as  quickly  same  2  spores  be  times  the  autotrophic  resting  example,  organisms  utilizable  actively  the were  i s safe to conclude  lysis  form  germination estuaries  It  primary  undergo  during  i m p a c t and  growing or  not  the  patterns  low  1 and  different  vessels with  a verylong  parts  both  phytoplankton.  being  growth  salinity  can  residence  planktonic  bacterial  as  most p a r t i a l l y  phytoplankton  the  10°/oo.  species as,  2,  both  not  than  experimental  biomass v e r y  the  in  can  may  they  have  suggests  to  results  they  Fresh  due  1 and  of  composition  that  the  phytoplankton  instead contents,  was  part  gradient  degradation  to species  obvious  only in  the  Comparing  becomes  different.  s e a w a t e r s y s t e m , even t h o u g h  after  systems.  it  appeared  d i d i n the  maximal  have  control,  time  they  This  most f r e s h bacteria)  t h e n go  through  to the  pathway. for  heterotrophic  components  for  two  different  58 salinity are  gradients  mainly  organic  controlled  activity  and  to  two  by  heterotrophic  as  that  a close  i n intermediate s a l i n i t i e s  The  or  changing  are  and  very  salinity)  ecological  system,  containing  the  system.  This  decline  of down  compared  in  the  system  bacteria  favours  energy  and  (Fig.  shifted 13,14).  to This  materials  vessels,  separation  of  the  phytoplankton  materials  supply  ( i . e . econiche)  input  (system  energy).  is  systems ( i . e . a  homogenous  a prime state  growth  state fora  with  a  bloom s u g g e s t i n g a  flowing  systems,  through  bacteria  the  d i d not  e x p o n e n t i a l growth as they b u t t h e maximum  bacterial  t h e d e g r a d a t i o n phase o f t h e p h y t o p l a n k t o n  provides  largest  are  phytoplankton  ecological  systems ( i . e .  I t i s a stable  In t h e c o n t r o l  experimental  the  controlled  during a phytoplankton  pathway.  the  of  condition  energy.  growing  1984).  systems  a co-increase with phytoplankton  biomass  The  T h i s may be  usually  experimental  with the  which  condition  heterotrophic  in  Controlled  lowest  of  (Spies,  arrangement  different  salinity).  in  of  microflagellates.  because f r e s h water b a c t e r i a  development  components  did  rates  s u c c e s s i o n i n g r o w t h between f r e s h w a t e r and s e a  better  show  utilizable  and  o f f r e s h water p h y t o p l a n k t o n .  bacteria  slow  populations  1) by  concentrations  water  stable  bacteria  parameters:  (both  2)  that  o f t h e h e t e r o t r o p h i c component may n o t be a s i n f l u e n c e d  salinity  due  by  substrates  production)  by  seems t o s u g g e s t  of l i g h t  energy  with  two  components  the  largest  i n t h e system.  (external  time  space  and  This creates the  energy) i n t o  However, i n e x p e r i m e n t a l  over  vessels,  ecosystem  the s a l i n i t y  59 gradient  creates  Phytoplankton This  left  heterogeneous  g r o w t h was  some  bacteria.  a  As  at  result, These  two  physical  environmental  response  of  an  the  conditions  ecosystem  to  the  u s e d by  b a c t e r i a showed  different  over  some p a r t o f  e c o s y s t e m s p a c e w h i c h was  a  phytoplankton.  inhibited  condition  gradient.  heterotrophic  a coevolution  s t r u c t u r e s under may  achieve  reflect the  time.  with  different  the  functional  largest  ecosystem  efficiency. The  ecological  phytoplankton gradient  (or  ecological can  not  space the  significance  and a  heterotrophic  a  heterotrophic what  is  more o r g a n i c  high  should  So,  the  than  i n any  components  input  i n the  may  kind  gradient.  behaviour the  (e.g.  than  i n any  of  aquatic  substances) i t . This  i n which  there  environment.  to e x p l o i t t h i s  component must p l a y  energy  The  source.  a more i m p o r t a n t  environment.  an  phytoplankton  organic  aquatic  salinity  suggests  successfully utilize  other  of  However, i f t h i s  head o f many e s t u a r i e s  have a s t r a t e g y  heterotrophic other  on  e c o s y s t e m s . The  system energy  happens a t t h e  ecosystem  different  components  of estuarine  occupy a c e r t a i n space  is  the  heterogeneous c o n d i t i o n a l g r a d i e n t )  evolution  contains  of  role  60 4.2  Experiment As  was  autotrophic group if  Stage shown  the  light  and  permits was  should  intensity  to  gradient  components  dilution water  components  are  below  gradient  In  exists  the  the  it in  up  experiments  part  gradient,  1,  experimental simulate  vessels  the in  population  allowed  provided  sedimental  The  experiment of marine The  stage  ecological  original  gradient  at  on  the  surface  II  a  marine  constant  layer  sea  during  entrainment  by  Simultaneously, water  over  also designed  a  time  salinity and  in three  over  parts.  gradient designed  w i t h an  water  increasing  was  water  to the euphotic l a y e r  (volume,  partially  by  input  planktonic  were  sea  a  sea  Thus  p r o c e s s w h i c h happens t o t h e  surface  w h i c h was  original  the  (determined  the  dark.  Increasing s a l i n i t y  In  water  p r o c e s s , marine ecosystem  is  the  energy  in estuaries  gradient.  along  brought  where  d i s t a n c e . The  go  components  mixing.  chain),  behaviour  salinity  simulating  ecological  estuarine  the  a temporal  rate  fresh  coupled with estuarine c i r c u l a t i o n )  observe  ecological  occurs  food  the growth o f the p h y t o p l a n k t o n .  on  4.21  phytoplankton  p l a y a dominant r o l e  components  from  a  based  r e s i d e n c e time  designed  I , the  the major f u n c t i o n a l a u t o t r o p h i c  remain  phytoplankton  phytoplankton  stage  g r a d i e n t i n the e s t u a r i n e ecosystem.  systems  (e.g.  loading  experimental are not  the s a l i n i t y  estuarine  that  in  components  on  pathway  II  was  61)  from  salinity mixed  to  flow  increasing  pumped  into  a covered  gradient  salinity  reservoir  to  time  which  water  seed  i n t o a body o f f i l t e r e d  fresh  estuary.  A  over  lighted  sea  « 0.20 o c  2 0 ^ o *  *  U 0.15-1 1  /  a  0.10-1  C  H5  Oi  10  u.  J k.  0.05] OjOO  \  •50  J z  ^  6  8  m  10  12  Days Figure  °I  15. The d e v e l o p m e n t p a t t e r n o f p h y t o p l a n k t o n and c o n c e n t r a t i o n o f n u t r i e n t s i n s t a g e I I p a r t 1 on an i n c r e a s i n g s a l i n i t y g r a d i e n t d u r i n g Aug. 1985.  0.0  Figure  16a. The d e v e l o p m e n t p a t t e r n o f t h e a u t o t r o p h i c component and c o n c e n t r a t i o n o f n u t r i e n t s i n s t a g e p a r t 1 i n an i n c r e a s i n g s a l i n i t y g r a d i e n t d u r i n g A p r i l 1985.  II  o>  5  Figure  —T"  10 Days  —i  15  16b. The d e v e l o p m e n t p a t t e r n o f NH 4 i n s t a g e I I p a r t 1 i n an i n c r e a s i n g s a l i n i t y g r a d i e n t d u r i n g A p r i l , 1985. +  20  64 water rate  in (  an D  increasing salinity  =  0.16  autotrophic indicated  / day  and in  ) ( F i g . 4)  nutrient  Fig.  15.  (NO~3  During  NO~2>  first  autotrophic  organisms  elimination  of  This  c h l o r o p h y l l a peak d e c r e a s e d  sea  water.  charge,  and  dilution  phase  until  inhibition  5  with  effect  populations  and  as  dilution  to  data  the of  inhibited an  The at  By  within  optimal  gradient  the no  longer  the  to  change  15°/oo).  Figure  a  increasing of  As  the  the  bacteria a  and not  This prolonged  lag  a t t r i b u t e d to  the  germination  of  seed  growing p o p u l a t i o n  as  well  shows t h e at  log  transformed  different  salinity  was  constant apparently  increased,  increased  salinity  by  toward day  5.  exponentially  gradient  spectrum  was  f o r marine phytoplankton  growth,  the  a h e t e r o g e n e o u s c o n d i t i o n so  the  sustained  to a phytoplankton  beginning,  surface  g r o w t h r a t e became f a s t e r stock  inflow  did  be  the  the  process:  in  low  marine phytoplankton  standing  range  18  the  work i n t h i s  a  on  of  filtration.  Chlorophyll  rate.  time,  f r e s h water  because  containing  growth of the  salinity.  this  whole system s h i f t e d At  the  a  thereafter.  may  salinity  c o n d i t i o n , the  chlorophyll  due  sea water  low  growth  low  optimal  The  on  high  to zero with  " c o n t r o l v e s s e l s " can  chlorophyll  salinities.  mechanisms  (salinity  of  days, the  two  grew  the are  some s m a l l  which  concentrations.  day  compared  by  of  of  components  z o o f l a g e l l a t e s during  sedimentation  nanoflagellate  increase  um)  by  Several  stress,  result  (<0.8  grazing  osmotic  high  a  +  the  at a c e r t a i n d i l u t i o n  evolution pattern  a  of  was  The  chlorophyll  small  value  gradient  (Fig.  17)  priority  bacteria  system.  had  a high  density  Figure  17. The d e v e l o p m e n t p a t t e r n o f b a c t e r i a and nanozooflagellates i n stage I I , p a r t 1 with i n c r e a s i n g s a l i n i t y g r a d i e n t d u r i n g Aug., 1985.  66  Figure  18. D e v e l o p e m e n t o f t h e a u t o t r o p h i c component i n a subsample c u l t u r e s e r i e s a t t e m p o r a l l y constant s a l i n i t y i n s t a g e I I , p a r t 1 d u r i n g Aug. 1985.  which  may  high  be  c h l o r o p h y l l a values  density  reached  phytoplankton appeared  at  growth  day  4,  the  effect  of  the  inhibition controlled  of  until  6, in  bacterial also  the  of  increase  release  in of  before  period  before  The maximum  density  until  t h e end o f t h e  a full  Flagellates  the f i r s t  demand.  population  To meet t h i s  a great in  increase the from  the be  in bacterial  density  nanoflagellate organic  products  period. 7  day 2  exponential great should  indicating a The  second  t o t h e end o f t h e  o f b a c t e r i a by  products  released  nanozooflagellate  i s indirectly  abundance. by  from  indicated a  The i n c r e a s e  by t h e  bloom as  the lack of c o r r e l a t i o n of  photosynthetic  offset  ( F i g . 17)  demand, b a c t e r i a  day  for  The  i n biomass,  first  by t h e  because o f the  growing  peak was r e a c h e d .  responsible  of  ina l l  The l o w e r d e n s i t y o f  started  phytoplankton growth.  may  3) a s  phytoplankton  a t the beginning  microflagellate  be  6 (see Table  (Fig. 13).  microflagellates  with  phytoplankton  increase  first  decreased  day  expected  activity  may  utilization  1981,  when  bacterial  experiment  The  the  t h e p a t t e r n may be e x p l a i n a b l e  experiment  have u n d e r g o n e  bacteria  i t declined. Bacterial  (by d a y 6 ) .  after  effect  is  biomass  increase  a s i n d i c a t e d by  microflagellate evolution pattern  filtration.  day  increase  during  slowly  But  nanozooflagellates  high  then  experiments.  the  A f t e r day 2  growing  phytoplankton  covariation and  .  cyanobacteria  B a c t e r i a d i d n o t show a c l o s e c o r r e l a t i o n w i t h t h e  of  other  of high  i t s peak  started  experiment.  in  composed  phytoplankton  grazing.  reflected  The d i e l  by  i n the  and p e r i o d i c  (Burney e t a l . ,  1982; E b e r l e i n , 1983; Hammer, 1983; K a p l a n , 1982;  Sournia,  68  Table  5. C o r r e l a t i o n m a t r i x o b t a i n e d f r o m t h e 6 v a r i a b l e s X 12 o b s e r v a t i o n s i n e x p e r i m e n t s t a g e I I , p a r t 1 d u r i n g Aug., 1985, between t i m e ( d a y ) , F l u o r e s c e n c e ( f l u o ) , N0~3, F l a g e l l a t e s ( f i g ) , B a c t e r i a ( b a c ) , and s a l i n i t y . P r o b a b i l i t y : * p<0.1, ** p < 0.05, *** p < 0.02, **** p <0.01.  I DAY FLUO N03 FLG BACT SAL  Table  1  | 1.000 0.539* -0.938**** 0.500* -0.381 0.983****  i  FLUO  1.000 -0.553* -0.230 -0.409 0.380  J  I  N03  j  FLG  j  1.000 -0.417 0.411 -0.900****  BACT  1 1  j  1.000 0.249 0.610**  1 .000 -0 .319  6. C o r r e l a t i o n m a t r i x o b t a i n e d f r o m t h e 6 v a r i a b l e s X 12 o b s e r v a t i o n s i n e x p e r i m e n t s t a g e I I , p a r t 2 d u r i n g Aug., 1985, between t i m e ( d a y ) , F l u o r e s c e n c e ( f l u o ) , N0~3, F l a g e l l a t e s ( f i g ) , B a c t e r i a ( b a c ) , and s a l i n i t y . P r o b a b i l i t y : * p<0.1, ** p < 0.05, *** p < 0.02, **** p <0.01.  DAY DAY FLUO NO 3 FLG BACT SAL  DAY  1.000 0.857**** -0.934**** -0.495* -0.177 -0.983****  FLUO  N03  FLG  1.000 -0.816**** -0.264 0.136 -0.821****  1.000 0.654*** 0.018 0.971****  1 .000 0 .067 0 .630**  BACT  1.000 0.166  69 1974)  may  limited may  create  by  situations  substrate,  s o t h e e x p l o i t a t i o n by  show a n e t r e d u c t i o n Table  the  correlation  variables;  bacteria  did  not  coefficient  with  In  part  vessels a  fluorescence  Decreasing  salinity  2,  and  decreasing  ecological development phase,  one  day  the  perturbation  compared  lag  germination  may  of  salinity  range  relative  to at  the day  its  peak  by  the  phytoplankton  s t r e s s • and standing turn  may  fresh water),  filtered  f r e s h water. T h i s  resulted in  over  and  with  decreasing  growth  salinity  was  may  salinity  reported  very  introduced have  some  ) , and  f o r many  be s l o w e r  growth  quickly  the  phase  and r e a c h e d  by day 6 s t o p p e d  decline  of  of the c e l l s .  activity.  nutrients the  1  ( F i g . 19) and t h e o s m o t i c  caused  caused  -  a t a lower  may  The e x p o n e n t i a l  increased have  The  a l s o a f f e c t the  the beginning  t h r o u g h t h e d e a t h and l y s i s  may  may  The o p t i m a l  at  rate.  marine  The e x t r a  (D = 0.16 d  The d e p l e t i o n o f n i t r o g e n  biomass  the  the " c o n t r o l l e d v e s s e l s " .  have s p e e d e d up t h e b a c t e r i a l  which  for  ( F i g . 19) shows a two day  Biomass i n c r e a s e d  dilution  water  time  f r e s h water b a c t e r i a .  phytoplankton  dilution  d a y 6.  stock  experimental  gradient  component  the  3.  correlation  in  i n c l u d i n g Skeletonema  so  the  I experiment.  s e a w a t e r was h e l d  seed p o p u l a t i o n s .  species,  any s i g n i f i c a n t  have been due t o d i l u t i o n of  seawater  started  with  of marine  lag  between  gradient  salinity  microbial  show  coefficient  as i n s t a g e  the o r i g i n a l  diluted  nanozooflagellates  effect.  shows  4.22  5  i n which b a c t e r i a a r e t e m p o r a r i l y  chl a This i n  The i n f l o w o f  (17.5 u g - a t N t o 6 1 o f  prolonged  plateau  phase  Figure  19. The d e v e l o p m e n t pattern of the a u t o t r o p h i c component and c o n c e n t r a t i o n o f n u t r i e n t s i n s t a g e I I , p a r t 2 ona d e c r e a s i n g s a l i n i t y g r a d i e n t d u r i n g Aug. 1985. o  Days Pigure  +  20b. The d e v e l o p m e n t p a t t e r n o f NH 4 i n s t a g e I I , p a r t 2 on a d e c r e a s i n g s a l i n i t y g r a d i e n t d u r i n g A p r i l , 1985.  73 observed  during  decline at  rate  least  of  the  f l u x was  April,  Aug.  1985  compared w i t h  part  nutrient in  the  experiment  the  b i o m a s s was  i n a chemostat i n the  f r e s h water c o n t a i n e d  nutrients  s t o c k may  sustained  a  temporally  balanced  state. This  implies that,  of  an  estuary,  certain  conditions  may  exist  chemostat environment which p r o v i d e s  larval  stock  feeding  resulting  i n a high  assimilate a  b i o m a s s ; t h i s may  and  Heterotrophic  With  l a r g e biomass s t a n d i n g  may  promote the  bacteria  inorganic  sufficient  have  carbon  a  of  longer  (Parsons very  organic the  autotrophs  nutrients.  Most  the  organic  products  photosynthesis nutrients organic by  are  f o r the carbon  Wolter,  inorganic reaction and  1982).  This  very  and  in turn  rate of m a t e r i a l s  evolution tightly  of  coupled  nitrogen with  can  inorganic during  nitrogenous  great  amount  q u i c k l y be  H a g s t r o m , 1979;  s p e e d s up  1970).  released  b a c t e r i a need  and  to  bacteria  favours  Martin,  This  bacterial  of  used  the competition  death of phytoplankton. growth  ability  for  The  of  efficiency  Painter,  p l a t e a u p h a s e may  (Larson  the  exponential  i n c r e a s e s the The  first  bacteria  nutrients after  so  s y n t h e s i s o f amino a c i d s .  r e l e a s e d a t the  heterotrophic  1980;  carbohydrates  in a  e t a l . , 1984).  carbon  with  as  zooplankton  strong  compete  mouth  periods  transport  successfully  of  i n the  briefly  n u t r i e n t s ( N i c h o l a s , 1963; source  be  b e n e f i t the  secondary production  out  much h i g h e r  in  standing  slow  experiment c a r r i e d  20a).  high  the  s t a t e , i f the  (Fig.  natural  The  but  "control vessels" indicates that  l a r g e enough as  i n which  ( F i g . 19),  for  chain  activity  recycling. (NO~3  +  NO~2)  was  not  the growth of p h y t o p l a n k t o n  at and  74 with  the  larger this  dilution.  than period  but  this  the  dilution  may  have  uptake  depletion  of  decline  water,  major  especially been  borosilicate  and  or  were  spp.,  was  day  15°/oo  results  of  6.  By  phytoplankton at water.  Skeletonema  from  low  of  spp.,  spp., but  day  6,  Ditylum spp.,  a l l were  costatum,  rate  which  rare. became  T h i s may have  selection  e v e r y day and c u l t u r e d i n series  state.  indicate  different  salinities,  is  to  e f f e c t s as  (1978).  i n order  areas  i n 25 x 150 mm to detect  This i s consistent with the t h a t most c o a s t a l of the world  1982;  a common e u r y h a l i n e  seems t o o c c u r a t s a l i n i t i e s  that the  had an o p t i m a l s a l i n i t y o f  Qasim  planktonic of  species of  show  (15 - 2 5 ° / o o ) i . e . l o w e r  1984,  the  under a c o n s t a n t  F i g u r e 22 i n d i c a t e s  phytoplankton  which  (Brand,  due  i n the experimental  Thalassiosira  i n our experiments.  others  by t h e  The c o n t i n u e d  obviously  of these phytoplankton  i n a batch c u l t u r e  about  growth  to  stopped  sources.  was  Skeletonema  tubes  germination  assemblage  sea  3  Asterionella  taken  culture  natural  growth  day  be  s p e c i e s were p r e s e n t  shown by H a r r i s o n and D a v i s  salinity  carbon  c o m p e t i t i o n and d i l u t i o n  Subsamples  was  _  species  to  N0~2>  growth d u r i n g  dominant a t t h e end o f t h e e x p e r i m e n t .  due  growth  +  (N0~3 + N0 2> were d e p l e t e d .  as N i t z s c h i a spp.  to  concentration  phytoplankton  Chaetoceros  (N0~3  The b a c t e r i a l  seemed  organic  from  nutrients  such  The  growth  of  the decline of nitrogen nutrients  bacteria  nitrogen  phytoplankton  Some  by  decline  effect.  caused  available  in  nitrogenous  The  et  maximum  than  normal  al.,  1972).  diatom,  optimal  (15-25%) ( B r a a r u d , 1951;  o  o o CO  Figure  21. The d e v e l o p m e n t p a t t e r n o f b a c t e r i a and nanozooflagellates i n stage I I , part 2 on a d e c r e a s i n g s a l i n i t y g r a d i e n t d u r i n g 1985.  Aug.,  77 P a a s c h e , 1975;  Brand, 1984),  Figure and  21  with  biomass  that  threshold biomass but  of of  grazing its  decreased  work These  similar  suggest  densities  The  lead  the  by  bacteria  data  through  occur  in  Skeletonema direct  studies  spp.  s t a r t e d growing  began  depends  antibacterial  exponential Spies*  ( F i g . 13,  when  Low do  activity  mediated  excretion  14). times  favourable  nanozooflagellate not  by  seem  external  (1979)  activity  was  1968).  to  be  confirmed an a  The  production  of  substances,  or  e t a l . (1974) of  this  stimulatory  algal  This  growth  observed during  bacteria result  inhibitory  phase.  metabolites  bacteria.  Bell  inhibition  has  and  the  inhibitory  and  exponential upon  through  of  1959,  costatum spp.  during  lowest  in bacteria.  Kogure  Vibrio  lower  at different  occur  period  stimulation  S.  the  a l s o shown i n  blooms.  this  (Sieburth,  that  and  basically  strongest  indicate  control vessel  inevitably  be  the  selective  evidence  Flavobacterium  consistent  b a c t e r i a was  biological  may  costatum.  Pseudomonas  quite obviously  r e l a t i o n s h i p between p h y t o p l a n k t o n and  substances,  a  and  phytoplankton  the  bacteria  nanoflagellate  may  different  during  for reduction  competition  reported  in  data.  heterotrophic  phytoplankton  phytoplankton  interaction  stimulative  of  results  to  important  the  my  nanozooflagellates. Bacterial  peak b e f o r e  when  mutual c o n t r o l of  specific  action  the  which  responsible  on  decrease  the  was This  results  that  conditions  an  and  higher  beginning  first  sharp  curve of  experiments.  sharply  This  (1984)  The  the  other  reached  growth.  at  i s consistent with  coevolution  nanozooflagellates.  bacterial  is  shows t h e  which  by  with  effect  on  effect  on  antibacterial  conditions. the  The  exponential  78 phase  of  algal  population  growth.  of  different  different  physiological  costatum).  The  exponential  growth  the  plateau  chl  a  phytoplankton are  mortality  occurs.  Considering  these  as  During  ambient  amino  species  my  is  agreement  the  during  found  found  a  the  during  that  of the  Vibrio-like  when p l a n k t o n  with  work done  by  e t a l . (1971). mode, t h e e x t r a c e l l u l a r  exponential et  only  al.,  carbon  the  grow a f t e r source  et  1982,  al.,  Bacteria  sources  nutritional  products  are  mostly  which  s o u r c e must t a k e up  nitrogen  which  growth  1984).  nitrogen  experiment,  attached  to waters,  different  bacteria  free-living  phytoplankton  natural  that  to  the beginning  in  a different  largely  after  Martin  and  with  were n o t  nitrogen. depleted.  the p l a t e a u phase  and  use  most l i k e l y  have  utilize  Bright,  1983).  This  also  mode may  dictate  different  groups.  In were  from  bacteria  (Skeletonema  species  This  during  (Amano  that  phytoplankton  later, particularly  Simidu  mineral  adapt  important  bacteria  acids  suggests  have  period.  period,  the  of  to  bacterial  the n u t r i t i o n a l  their  groups  different  (Eberlein,  this  However, no  was  phytoplankton  carbohydrates  states  the  i t s composition  phase of p h y t o p l a n k t o n  more  (1968) and  shift  generic  dominant  death  organisms  of  (1980) showed t h a t  d u r i n g t h e e v o l u t i o n may  succession  Sieburth  Martin  diatom  growing  while  during the  attached bacteria  e x p o n e n t i a l g r o w t h . The f r u s t l e s , and  free-living bacteria morphological  first  and  dominated  l a t t e r were  were b i g and  period  mostly  rod shaped.  attached bacteria  p r o p e r t i e s and  a l s o may  In may  require  different (Fukami in  nutritional et  a l . , 1985,  this  part  composition. decrease  the  and The  and  P  that  not  suppression  which  for  that  the  Kogure  influence of  organic  peaks  which  inhibitory may  be  density nutrient  in  substances  to  of  nutrient bacterial before  c o m p e t i t i o n was decline  inhibitory growth  shift  decreased  (1979) showed t h a t  bacterial  developed  potential  strain  bacterial  the  substrates  reflect this  another  bacterial  indicates  bloom.  two  excrete  stimulate  mechanism  did  dissolved  e x p e r i m e n t may  bacteria  to  depletion  direct  that  the  fact  phytoplankton N  of  P h y t o p l a n k t o n may  competitors  nutrient  of  1 9 8 3 , 1 9 8 1 ) . The  free-living  growth.  modes  was  before  the  effect not  not the  addition  and  of  concluded  a  result  of  competition. He  also  found  decreased light  the  inhibition  occur  and  the  phytoplankton there  must  was  between b a c t e r i a  may stage  a  and  give for  way  bloom, to  the  activity.  What  is  priority the  intensities be  due  which reduce the the  the  inhibitory  growth  of did  growth  gradient. of  the  effect  i n which the  salinity  to  of  Thus  interactions  phytoplankton. that  under the  c o n d i t i o n s of  heterotrophic  pathways  u t i l i z a t i o n of  mutually  autotrophic production.  autotrophic  T h i s may  I experiment by  light  more complex d y n a m i c s t a t e  T h i s phenomena i n d i c a t e s autotrophic  lower  e x p l a i n why  inhibited  exist  and  light limitation  This  in  higher  antibacterial  phytoplankton. not  that  Then t h e  are  an  s h u t down  required  to  nutrients  w h o l e s y s t e m becomes  an  system.  effect  of  this  interaction  on  microflagellates?  80 There a  i s no i m m e d i a t e answer t o t h i s  higher  trophic  affected. 6  The c o n s t a n t  i s quite different  bacterial the  depend on b a c t e r i a , t h e y  low l e v e l  from o t h e r  population  results  responses  of  gradient.  of nanozooflagellates experiments.  ( F i g . 21) d u r i n g  to  range  10  of  the  same  not  actively the  are at must be  u n t i l day  The d e c l i n e o f t h e  first  that  p e r i o d may  but  populations  may  they  prevent  have  a  wide  cultures  seed p o p u l a t i o n  function This  suggests  a t the higher  results  to  or germinate  occupy  occupy the  phytoplankton  low  adapted  extrapolating  heterotrophic  show t h a t  the  d i d n o t grow d u r i n g flasks.  They can portion  seawater p o p u l a t i o n s can spectrum. suggest  In other salinity  part  words, b a c t e r i a gradient,  of s a l i n i t y  while  gradient.  b a c t e r i a occupy t h e low s a l i n i t y  w a t e r a t t h e head  at  the  result  the  high  below  i t means t h a t t h e  the estuary, mouth  on t h e low  p e r i o d o f t h e e x p e r i m e n t and  part.  of  of  t o an e s t u a r y ,  of  this  that  first  last  part to  range  two p a r t s o f t h e e x p e r i m e n t s  the  the  salinity  i n t h e low s a l i n i t y  end o f s a l i n i t y  i n these  different  to a  c a n be i n h i b i t e d  Subsample  the  the quite  p e r i o d o f time as i n t h e e x p e r i m e n t a l  phytoplankton  field  populations  (<10°/oo).  bacteria  adapted  show  phytoplankton  salinity,  only  The  experiment  salinity  gradient.  flourish  this  seawater  salinity o/oo  in  Seawater  adaptations  of  and  s i n c e they  r e s t o r a t i o n of the nanozooflagellates. The  In  level  question,  and p h y t o p l a n k t o n of  observations  the  estuary.  (Albright,  standing This  stock  appears  greater  i s i n agreement w i t h  1983; B e l l  many  e t a l . , 1981; W r i g h t ,  81 1984,  1985).  Correlation coefficient expected  fresh of  the  appear The  between because  However, not  calculation  controls and  23a,b).  had  two  f l a s k s are quite  standing bottom due  water  two  culture  of  were  and  is  phytoplankton.  containing  The  evolution  pattern  i n t h e s e two  flasks  Thalassiosira  I used a s t r o n g  by h e t e r o t r o p h i c  contains  u g - a t N/1.  depletion  i n these  r e s u l t from the  Sea w a t e r  17.5  maximum  spp. i n sea water  The maxima o f b i o m a s s  difference.  original  of the c h l o r o p h y l l  d i f f e r e n t ; t h i s may  stock a f t e r nutrient  to degradation  This  and p h y t o p l a n k t o n d i d  similar  species  f r e s h water c o n t a i n s  though  salinity. the  vessels  respectively.  respectively.  concentration  (even  and  development  major  costatum  fresh  while  N0~3,  assemblages  and  N/1,  fluo,  of the experiment.  sea water  The  Skeletonema  nitrate  the  i n t h i s part  phytoplankton  were  of  6) shows a h i g h e r s i g n i f i c a n t  c o r r e l a t i o n between b a c t e r i a  water  (Fig.  time,  (Table  i s due  35 u g - a t  The d e c l i n e  to sinking  magnetic s t i r r e r )  bacteria  initial  to the  and  ( F i g . 13,14).  of  also  o  Q£ 0»0  i  0  Figure  i  i  i  i  5  i  I  i  I  i  10 D a y s  i  »  i  I  i  |5  23a. The d e v e l o p m e n t pattern of the a u t o t r o p h i c component i n a c o n t r o l l e d s y s t e m w i t h o r i g i n a l s e a w a t e r d u r i n g Aug. 1985.  '  20  0  3.0i  ;ence  25  20 u. >  \&  1.0 05'  rr 00 10  15  20  D a y s Figure  23b. The d e v e l o p m e n t p a t t e r n o f t h e a u t o t r o p h i c component i n a c o n t r o l l e d s y s t e m w i t h o r i g i n a l f r e s h w a t e r d u r i n g Aug. 1985.  00  84 Disscussion  There structure role  is surprisingly  of  f r e s h water e n t e r i n g the sea or  Some  phytoplankton al.,  work  in  has  estuaries  1968;  on  the  physiological  than  on  the  ecological  al.  (1969)  diluted almost that  sea  It  phytoplankton  some  normal  biomass  with The a  the  my  pass  i n an  estuarine  reported  on  in  water  °/oo)  can  survive  an  mostly  ,  on  small  study,  in  focus groups  in  can  not  estuarine  ( F i g . 5a, controlled  . Along  concluded  ) clearly  10a,  The  as  an  living  by  freshwater  11a  ) compared  ( F i g . 13,  was  this  demonstrated  function  systems  systems  of  structure.  ecosystem.  6a,  saline  estuarine  i s the e f f e c t  actively  or  gradient with  the whole ecosystem (stage I  in  freshwater  the s a l i n i t y what  Blanc  dead  diluted  autotrophic role  experimental  of s a l i n i t y  (Blanc  they  whether  of the freshwater phytoplankton  range  water  phytoplankton  were  matter  not  i n the  fresh  of these phytoplankton  through  If  in  very  production  component  fresh  p l a y an  phytoplankton  was  ecological  F o e s t e r , 1973), but  26  experiment  contribution  activity narrow  still  component  phytoplankton  -  phytoplankton  freshwater  autotrophic  that  important  they  in  of the  (1973) i n a more d e t a i l e d  functions? of  results  that  can Can  disfunction The  an  ecosystem  i n the e s t u a r i n e ecosystem.  phytoplankton  is  the  s a l i n e water environments  state  (8°/oo  Foester  freshwater  ecosystems.  and  role  concluded  water  dead.  waters.  been  W e t h e r e l l , 1961;  more  et  i n f o r m a t i o n on  o f the f r e s h water p h y t o p l a n k t o n  ecosystem.  et  little  14).  restricted  the whole s a l i n i t y  in  gradient,  85 most the  of  the freshwater  organic  change  the  a  component  became  high always  which  major  as  that  of  to  a  sea water  in  the population  sea  (Stage  I,  Part  may  gradient occur  at  system.  a  the  gradient grow  the in  niches  an  f r o m low t o  i s i n a condition of  dominance g r a d u a l l y r e a c h e s  of  condition  the  of  This  of  the  gradient  is  delayed  almost  i n an  gradient  separates were  the  inhibited  ( F i g . 1 5 ) , as t h e  s u i t a b l e range,  I f we c o n s i d e r water  i n the  i t s maximum  salinity  which c l e a r l y  salinity  biomass  but  gradient.  organisms i n  i s dominant  over time. Marine species  vessels,  element  gradient  g i v e a p i c t u r e o f what  q u i c k l y and t h e b i o m a s s i n c r e a s e s  salinity  i n t h e same  of freshwater  end o f t h e g r a d i e n t .  development  controlled  been  has  ( F i g . 5a, 6a; F i g . 1 5 , 1 6 a ) . A t t h e  moves c l o s e t o a s p e c i e s  very  maximum  This  end  i s probably  s t a t e . The t i m e s e r i e s o f  1) t o g e t h e r  estuary  heterogeneous  low  This  t h e sea water p o p u l a t i o n  salinity  phytoplankton at  a real  phytoplankton  1) and t h e s e a w a t e r o r g a n i s m s  I I , Part  level,  the high  creates  (Stage  in  phytoplankton estuarine  gradient.  phytoplankton  change  water  i n t h e f r e s h water  the s e a water  t i m e and s p a c e . So t h e e c o s y s t e m close  flow  ecosystem.  the evolution of the s a l i n i t y  being  structural  the heterotrophic  contribution  I I showed  evolution  into  the energy  pathway. T h e r e f o r e  biological  same  shifted  function i n a s a l i n i t y  the  over  the  stage  quickly  direction  system  the estuarine  Experiment  because  whole  heterotrophic  component o f  can  b i o m a s s was c o n v e r t e d  d e t r i t u s p o o l . T h i s change r e s u l t e d i n a  for  towards  phytoplankton  phytoplankton  e x p o n e n t i a l l y . The t h e same as i n t h e  growth phase r e s u l t s t h e same p r o c e s s  from  happening  m o v i n g s e a w a r d on t h e s u r f a c e  layer  86 of  an  estuary  the  components  in  pattern.  This  temporal e v o l u t i o n p a t t e r n  that  element  model  estuarine  ecosystems  The  that  fact  may  be  show  the  will  spatial  i t s maximum o f t e n a t some d i s t a n c e  estuary  i s an  The  dynamic p a t t e r n  experimental  systems  phytoplankton  and  because  the  between  living  process  is  biomass  N0~3  process  of  the  of  a  time  N0~3,  which development  of  nutrients  also  necessary shown  in  phytoplankton estuarine  are  the  biomass  with  spatial  along  an  be  along in  along  negative  easily  the  an  an  estuarine  which  an  estuarine  component  s c a l e by  estuary  should  covariance  i n t o the  living  and  bacteria).  negative  correlation  correlation as  between  a result  of  distribution  salinity  gradient  is  i n c r e a s i n g biomass  of  transition  from the  be  expected  growth  and  outflow,  the the  In  estuarine  Because  head o f  a standard  zone.  some  components.  continuous  of  i s , the  explained  seawater phytoplankton  the  i s t o be  decided  strong  major a u t o t r o p h i c  e v o l u t i o n of  coincided of  i s found  a  spatial  evolution  over time. T h i s k i n d of  works  ecosystems,  populations temporal  all  and can  phytoplankton  of  (phytoplankton a  the  the  n u t r i e n t s i n a l l our  incorporating material  show  gradient  mass.  n u t r i e n t s . That  components  and  with  dynamics. T h i s  and  why  pattern.  a salinity  function  relationship  calculations  Chl  and  bacterial  components  a  water  a  of  f r o m t h e mouth o f  of nitrogenous was  nutritional  Statistical between  explanation  e v o l v i n g p a t t e r n which c o i n c i d e s  b e c a u s e o f t h e movement o f t h e  distribution  distribution  i n c r e a s e of c h l a along  reaches  scale  the e c o l o g i c a l  become a s p a t i a l  a reasonable  this  of  the  estuary increase  phenomena  as  87 the  phytoplankton  conservative pattern. quite  nutrients  different  behaviour  as  a  is  t h e major  extremely  variable  nutrient  unicellular  time  may r e s u l t utilize  in  amino  (Adernson,  algae  +  and  as energy  which  may  and ammonium  experiments  take  a  testing  decrease  t o zero i n the f i r s t of  due  C h i a.  t o uptake  different. Bacteria  be g r a z e d by +  release It is  dynamics  of  There  from  NH 4 interesting +  NH 4  showed  components b u t n o t w i t h  experiment,  value  p e r i o d o f t h e experiment, was no i n d i c a t i o n (this  In t h e  the concentration of  i t s initial  by p h y t o p l a n k t o n  this  1 9 7 2 ) . The  g r a d i e n t s ( F i g . 5b,6b, 16b a n d 2 0 b ) .  components  shows  up  i s very  1985).  system  is  i n the presence  s o u r c e and then  the  their  1967 M c C a r t h y ,  dynamic p a t t e r n s .  and C a r o n  salinity  was  known t h a t  directly  different  increase  (Dugdale,  N0~3  quite different  patterns with d i f f e r e n t  freshwater  NH 4  assimilation  preferentially  NH 4  acid  i n my  +  utilize  of nitrate  nitrogen  and space  different  close  can  organic nitrogen, although  I t i swell  1985; Goldman  that  +  bacteria  total  time  of  microflagellates  NH 4  o f ammonimum  ( B l a s c o a n d Conway, 1982; M c C a r t h y & E p p l e y ,  generation  can  to  Walsh e t a l . , 1980). +  The d y n a m i c s  has a  o f many p r o c e s s e s w h i c h happen i n and  to  in  spatial  p r o c e s s by w h i c h t h e s e two components  nitrogen  contribution  This  pattern.  biological  inorganic  NH 4,  corresponding  s o u r c e . The a s s i m i l a t i o n  relative  of  a  o f ammonimum i n t h e s y s t e m  Phytoplankton  nitrogen  1972;  have  i s a combination  ecosystem.  covert  also  s p a t i a l p a t t e r n and t h e p r i m a r y  However, t h e d y n a m i c s  concentration the  distribution  that  this  t o a value before the decrease  i s most c l e a r  i n the  o r g a n i c based system ( h i g h I n t e r n a l energy)  thermodynamlcally unstable  I  heterotrophic p r i o r i t y systems s t a t e  A J nutrients i <  I I <I X  external (light) energy  x i nutrients •photosynthetl process.  « c  external (heat) energy  A  shortwave radiation  ±=  6  e t e r o t r d pp h l c process  i (Concurrent' ! state ' ) , , T , nutrients | 1  respiration longwave radiation  -  , , {nutrients J autotrophic p r i o r i t y systems s t a t e  Cheriaodynamlcally stable  « ( •  I n o r g a n i c based system ! (low I n t e r n a l energy) I t  ground  state  L energy  flow oo 00  Figure  24.  The e n e r g e t i c dynamic s t a t e the energy l e v e l field.  o f an e c o s y s t e m  in  89  experiment of  N0~3  still  during over  showed +  NH 4  (  which  grow  in  at  extra  the  of  NH 4  first  nutrient the +  of  NH 4  salinity  uptake  that  of  may  stage  result  an  as I  +  The  from  NH 4  the  higher  from  initial  peaks  detrital  of the  may  indicate  differences i n the  ratio of  difference i n the  i n e i t h e r water  w a t e r s had a l r e a d y other  nature  ( i f we assume reached  factors).  of  fresh  stress  w h i c h may c a u s e t h e r e l e a s e o f  the  aphotic  In  w a t e r and s e a w a t e r  f r e s h water organisms. T h i s  of  cell  regardless  inflow  system,  the  increasing  some  n u t r i e n t s may s t i m u l a t e  in  had  by  organic  bacteria  but  patterns  determined  because  amino a c i d s . I n t h e  c o m p o n e n t s . T h e s e two d i f f e r e n t  substrate  carbohydrate  need an  materials  osmotic  seawater  and t h u s t h e y  organic  organic  the  the bacteria  e x p e r i m e n t were most  degraded,  populations  biomass  experiment  in  low  t h e b a c t e r i a i n both t h e i n i t i a l  in  bacteria  b i o t a , t h e dynamics o f t h e  phytoplankton  result  bacterial  maximum  The h e t e r o t r o p h i c  synthesize  i n s e a w a t e r and f r e s h w a t e r o r a the  t h e NH 4  suggest that  seawater  gradients.  heterotrophic  +  may  was  +  but  period of the  to  decrease  b i o m a s s w h i c h may p o s s i b l y  are quite  phytoplankton  mineralization  by  of  the  involving  after  different  C/N  F i g . 7 ) . This  nitrogenous  peaks  increase of  period  expense a t c a r b o h y d r a t e ,  concentration  the  an  the  experiments  experimental  a d e c l i n e over time.  utilize  likely  ( F i g . 5 b , 6 b ) ) . T h e r e was no  t h e whole  showed  production  April  addition of  t h e g r o w t h o f b a c t e r i a . However, the  water  may  long  history  of  contain  u t i l i z a t i o n by  z o n e . The s e a w a t e r p o p u l a t i o n  concentration  little  had a  w h i c h may have r e s u l t e d i n  90 rapid  uptake  of  +  NH  before  4  the  start  of  exponential  growth. The  high  metabolic  heterotrophic to  go  back  components to  the  energy  dominated  by  potential  bioenergy  which  From  my  behaviour  so  be  gradient.  In  (tide  average)  fresh  water  real  there  is  to  s a l i n i t y  euryhaline  most  of  and  ranges  contribution  compared  The  seawater  can  be  along  environment.  The  estuary, only  the So  provides most  an  this  the are  the  important  one  gradient  a  tendancy  is  the  of  when  the  system  the  is  largest  Therefore  the  low energy  containing  whole  into  feature  a  moves  of  s a l i n i t y  To  f i l t e r ;  i t  they  material. function  gradient,  different  more  gradient.  as  but  population  them  c i r c u l a t i o n  a  their  s a l i n i t i e s .  favourable  the  s a l i n i t y as  is  very  However,  the  close  their  actively  as  pass Some at their  behaviour.  to  the  seawater  direction  organic  seawater  function  in  that  and  phytoplankton  s a l i n i t y  have  show  works  keep  the  gradient  more  and  component  increasing  may  to  they  that  clearly  with  converted  low  a  has  having  different  freshwater  of  at  on  results  t o t a l l y  phytoplankton  inhibited  capacity.  autotrophic  phytoplankton  intermediate  level, and  based  system  system,  lowest  is  the  components,  out  the  mineralization  stable.  is  the  f i l t e r s  gradually  more  freshwater  component  movement  i t  experiment, of  that  substances  that  may  fast  p r i o r i t y  its  carrying  autotrophic  high  at  inorganic  adjusts  component  is  and  suggest  autotrophic  internal  system  rate  low. They  continual  the  optimal outflow  conditions. an  ecosystem  is  the  dynamics  91 of  the  phytoplankton  effect  on  closely  the  second  in  based  these  on  as  gradients  as  did  our  that  Because  dependent There  different  system state amount  both  Their  coupled  one  are  two  The  which  most  are the the  bacteria  are  activity of  and  organic  d i f f e r e n t foundations  decide  systems  different  dynamics with  and  energy  is  energy  shown  F o r two  different  be due  salinity biomass salinity  t o the presense of  freshwater. show a  gradient  components, range o f  that  And  tolerance  also  either  t o a range o f  a r e much more  the  the  is  by t h e s a l i n i t y  amount  and  quality  the s t a t e of the a u t o t r o p h i c  c a n have two  i t  d i d n o t show a c r i t i c a l  seawater  considers  which  their  heterotrophic  or freshwater b a c t e r i a  If  so  p h y t o p l a n k t o n . T h i s may  and  radical  p o o l and  on t h e a v a i l a b i l i t y  of phytoplankton.  seawater  substrates  ecosystem.  heterotrophic  i s not a f f e c t e d  in  tightly  have a  substance  transformer,  bacteria  salinities.  the  organic  experimental  of b a c t e r i a  the  must  ecosystem.  h a v i n g two  growth  of  components  productivity  much  the  water. two  a l l  bacterial  and  the  i n the In  as  energy  is primarily  substance  routes  pool,  which  components t o p h y t o p l a n k t o n  components.  level  production  structure  ecological  nutrient  heterotrophic a  whole  related  inorganic  production  energy states,  thermodynamically  organic  component.  state  one  of  of a ecosystem,  i s a high  unstable.  internal  There  is a  the  energy great  of o r g a n i c  substances i n the system. T h i s  system tends t o  internal  energy out of the system which  goes  release inorganic  substance  based  system;  this  i s a low  back  internal  t o an energy  state  and  is  major  role  i n t h i s energy r e l e a s e  high  uptake  inorganic which  limited  properties natural  of  by  the  internal  This  the  state  ecosystem, level  system)  This  i t  heterotrophic processes, potential uphill  As  is  means  will  may  be  level  light  the  mainly  The  when t h e  physiological  energy  energy  with  state  back  to  ground  bacteria).  stable.  to a certain of the  the l i m i t i n g system  (living  o f the ecosystem state  Between  through these  processes are downhill processes; than a u t o t r o p h i c  because  they  and e n e r g e t i c  privilege  of  a two  their  p r o c e s s e s which a r e  needs e x t e r n a l  i n a diagram  the  into the  capacity by  from o u t s i d e  isa  through  increases  controlled  their  process i s  flows  a high energy s t a t e input  component  the bacteria  The o t h e r  greater  i s summarized  (u)  i s thermodynamically  o f the system  (mainly  heterotrophic  physiological  bacteria  go  growth r a t e  heterotrophic  external  the  that  process  processes,  idea  The  by  always needs energy  otherwise  This  excited  capacity  and  the system. A l s o  i s d e t e r m i n e d by t h e e n e r g y c a r r y i n g  this  nutrients.  which  a  substances  the autotrophic  i n t e r n a l energy.  play  have a v e r y  actually are consistent  state  process.  high  source.  The b a c t e r i a  the energy s t a t e  which  system;  role.  organic  to a certain  energy  bacteria  be  in  Bacteria  Bacteria  as a  over  high  the  energy  photosynthetic  is  system's  can  as w e l l  decreases  by  ecological  limited low  state  both  compete  state  stable.  process.  for  m  nutrients  energy  energy  become  (V )  successfully  the  system  thermodynamically  rate  mineral  can  when  more  energy  support.  (Fig. 24).  properties  utilizing  of bacteria  i t s energy  provide  source  as  quickly  and  ecological quickly  role  to  reflected When  as  a  i n an stable  in  the  organic  immediatly into  a  go  that  at  state  nature  of  become  limiting,  no  the  increase  ability  i s no be  be  effectively  utilized,  stable  state),  can  developed  bacterial  a  (Azam  low  K  m  (since  this  et  to  source.  concentration Morever,  due  food  have  nor  K  al.,  m  the  can  their Implicit  for the  the  input.  experiments  inwhich  substrate  (Parsons  al.,  1977,1981)  and  can  organic  the  in this  bacteria  Further,  not  go  ever  can back  become  l i m i t a t i o n on i s that at  low  of  added  the  bacteria nutrient  in seawateris  changing  been  to  priority  substrates  system c o u l d  material  a  microflagellates.  a l . , 1 9 6 7 , 1 9 6 9 ) . In  the  means  back t o  substances.  in t h i s theory  has  nature  system.  Bacteria  feeding  with  Vaccaro et  the go  go  they  (phytoplankton  concentration  change  of  not  but  This so  microzooflagellates  organic DOM  bacteria. biomass  organic  own  do  whole system would tend  energy  (so  1981;  et  The  amount o f  the  bacteria  ecosystem  also  population.  t h i s storage  of  between b a c t e r i a  neither  back  phytoplankton did,  l e t an  l i m i t e d by  ever  may  that  s y s t e m t o go  i n energy s t a t e  to  further  equilibrium  its  bacterial  storage  possible.  means  fully  as  certain  This  to  the  i n t e r n a l energy s t a t e  should  the  accomplish  f a s t r e s p o n s e mechanism i s  a  soon as  there  to  which suggests t h a t  the  low  if  is  to  as  a  systems)  there  The  depletion  keep  has  state  system  dormant  stage  respond  process  stay  possible  a high background c o n c e n t r a t i o n  quickly  stable  state.  through  bacteria  This  as  ecosystem, allowing  nutrients  dormant  provides  completely  low).  conditions a number  to  of  seawater  showed a v e r y  high  K.  This  m  suggests  population  from  concentration of  bacteria  can  balance  was  not  growth the  the  at  a  cell  growth The  of  numbers  the  organic  nutrients  upper  limitation which  is  than  a  growth  of  bacteria by  The and some  equilibrium  being  a  function  bacteria  i n the  only  After  more an  of  of  state  by  under  organic could  substance the  living  not  limit  the  and  can  not  Bacterial  go  lower d e n s i t y  of  into  a  limit,  microflagellate  exploiter  ( W r i g h t , 1978, is  bacterial  (e.g.  bacteria  the  that  availability  of m i c r o f l a g e l l a t e s  ecosystem.  of  offset  between o r g a n i c  which  growth  where b a c t e r i a l  bacteria  of  mechanism  input  a function  threshold  equilibrium  the  Some work i n d i c a t e s  the  are  with  experiment,  organic  may  decides  authors  my  bacterial density.  factor.  grazing  the  f l a g e l l a t e s can by  up  microflagellates  i s decided only  Microflagellates  suggested  of  the  limit  grazing  the  when t h e  is  substances of  In  the  substrate  zooflagellates  its physiological  grazing  limiter.  with  nutrients.  a space d e n s i t y  the  populations.  of  limit  state,  of  comsumption  bactivorous  upper  and  of  within  m  the  means t h a t  substrates.  microflagellates  organic  concentration  This  bacteria  K  always c a t c h  microflagellates  dormant). Therefore  control  can  g r o w t h . The  rate  the  depending  increased  of  regardless  of  shift  grazing  organic  always  i s l i m i t e d by  utilization  as  of  certain  abundance  may  high  substances.  phytoplankton  limited.  bacteria  very  l a r g e l y exceed the  by  not  to  organic  biomass  substances  bacteria  i t s growth r a t e  concentrition  bacterial  or  low so  addition  high  that  1982,  against  of  bacteria  substances,  doesn't  exist  1984). the  This  actual  95 The  properties  requirement to  the  provide  addition  several  fold  dynamics  of  absence stage  as  is  the  organic  pool  the  from  the mixing  initial  bacteria, of  labile other  which  in  the  organic  reflection  of the  experiments. period  The  In the  o f experiment  were  substance  both  to  source,  substance  o f water which  growth.  before  or  provide  could  the experiment limited  t h e n c a u s e some  a labile  source,  i t sutilization  which c o u l d  This  i n an o r g a n i c  must  addition  be some  i n one t y p e o f w a t e r b u t r i c h  I f we assume t h a t  factors  the  o f two k i n d s  the mixing process  organic  a  the f i r s t  may be s h o r t  waters  organic  increase  the increase of  indicate  other.  responding  ( W r i g h t , 1978, 1 9 8 3 ) .  actually  mutual b e n e f i t f o r the b a c t e r i a l  in  functional  n u t r i e n t s and i t c a n o f t e n  bacteria  1,  ecological  the a b i l i t y of q u i c k l y  phytoplankton during  micronutrients  an  a 24 h o u r p e r i o d  should  resulting  of  the  I, I I , part  bacteria  a  of organic  the  of  bacteria  bacteria  within  of  dynamics  of  be r e l e a s e d  state f o r  transformation  or, i f there  must  started  be l i m i t e d  by m i x i n g  i s some by some  t h e two  kinds  water. In  an  isolated  system, t h e o r g a n i c  autotrophic  component.  autotrophic  component  heterotrophic However,  in  phytoplankton from  the  increase  which  activity  is  depend  entirely  the  from t h e  supply  on  control of  the the  substrate.  e c o s y s t e m , t h e c o n t r o l o f b a c t e r i a by  perturbed  by  allochthonous  environment.  importance  bacteria  can  through  an e s t u a r i n e  terrestrial the  Therefore  s o u r c e must be  of  their  Therefore  organic  substance  t h e b a c t e r i a may  r o l e i n t h e e c o s y s t e m due t o  96  the  independent  much  bigger  energy on  role  flow  the  course  and  ratio  of  than  supply.  They w i l l  i n a autochthonous system  matter  of  substrate  recycling.  available  i n terms  Their quantitive role  organic  substances  and  play  a of  depends  the  mineral  nutrient. The 1.  e c o s y s t e m s t r u c t u r e may  Phytoplankton In  these  growth.  Bacteria The  substances  down.  and All  autotrophic part 2.  2 and  the  exudate  of  through  a l l "control and  limited  pathway  during  by  their  autotrophic  component c a n  system can  has  substrate  for  phytoplankton  of  released  on  bacteria is  material correlation  from  not  is  shut  provide  energy source,  i f the growth  use  The  obvious.  most  phytoplankton  f o r some o t h e r  phytoplankton  II  systems.  of  bacteria  calculations.  that  23a,b).  But  n o t make f u l l  show a c o e v o l u t i o n p a t t e r n .  of  in  extra  is restricted  may  experiment, stage  photosynthetic production.  the  so  n u t r i e n t p o o l goes t o  o r g a n i c n u t r i e n t must come f r o m r e l e a s e o f  component  indicated  populations  heterotrophic coevolution are  organic  i s that inorganic  bacterial  v e s s e l s " ( F i g . 13,14,  bacteria  by  of phytoplankton  i s shown i n o u r  autotrophic  bacteria  system  bacterial the  phytoplankton  limited  growth  possible mineral as  favour  less  the  quick  the  This  or  existing  energy flow  Phytoplankton  more  The  previous  the  conditions  foundation  productions  Provided of  the be  dominate.  any  material  may  forms:  systems.  systems,  substances.  inhibit  priority  show t h r e e  reason,  the and  system. so  inhibiting  the effect  However, t h e by  bacteria  These c l e a r l y  of  use is  show i n  our  experiment  gradient this 3.  stage  condition  case the  I  and  depressed  carbon budget  Heterotrophic p r i o r i t y The  system  I I ( F i g . 7,  at  the  showed  estuary  and  p o l l u t e d water.  by  phytoplankton is  source. bacteria with  under  the  material  goes  bacterial from  a  This  biomass  energy  substance  which  a  terms  within time  above an  population different  for  —  energy the  dissolved  to  The  Then t h e  priority  attempts  I f we  autotrophic (1977,  system  s t a t e and  input  state  phytoplankton the In  two my  to  consider  o f t h e same p h y s i c a l  groups.  Most o f t h e  o r g a n i c matter  energy  of the  provide bacteria  system.  a low e n e r g y  external  discussion  level,  energy  1981) occurs  through returns provide  to the  system  and more s t a b l e i n  structure.  ecosystem.  series  bacteria  state  phytoplankton  of ecosystem  The  through  i s a h o m e o s t a t i c mechanism w h i c h the  an  bacteria  m e t a b o l i c growth  and m i n e r a l i z a t i o n .  available is  in  of  in  o f an a l l o c h t h o n o u s e n e r g y  enough  dominate  eliminates  high  component  d e p r e s s e d , as shown by P a r s o n s  (1984).  and  of  control  then  and  rapidly  Then t h e  happens  t h e h e t e r o t r o p h i c pathway.  is  Spies  often  into  production  So i n  have a h i g h a l l o c h t h o n o u s  very strong  conditions to  salinity  c o n c u r r e n t pathways.  as  by t h e a d d i t i o n  case,  ability  such  autotrophic  overruled  In . t h i s  two  b e g i n n i n g may  addition,  supply  the  system.  substance  the  that  the p h y t o p l a n k t o n growth.  organic  some  12)  —  look at the this  s p a c e , we  c y c l e of energy  experiments  in a  have a s u c c e s s i o n  bacteria.  bacterial  interactions  However,  at  may  belong  maxmia most  of  of the to  autotrophic  98 component is,the  development  bacteria  different  from  previous to  that  autotrophic  so  They  energy  a l l  may  in  allow  not  inhibit  nutrients  depletion  energy  with  degraded  phytoplankton  necessary  biological  been  those  left  in  for  the  phytoplankton. material  elements,  phosphorus  such  The  the  system The  them  f i r s t to  which  compounds.  the  to  this  take  At  limited of  by  organic inorganic  group  this  into  time  organic  growth  of  exhausts  up.  at  There  They  l i k e l y  acids  its  living  period.  contains  amino  during  second  bacteria  They most  as  in  heterotrophic  period.  The  the  the  of  the  with  due  incorporated  capability.  from  group  causes  end.  likely  with  +  for  the  mineral  NH 4  release  growth.  nutrients  carrying  nutrients  organic  of  to  different  enough  have  in  bacteria  pool  normally  The  degradation  up  of  be  compete  this  quite  phytoplankton  are  and  the  are  or  may  grow  during  competitors  nitrogen  to  of may  system.  appear  components  not  natural  exponential  autotrophic  are  a  bacteria  take  group  with  are  most  interaction  growth  competitor  those  and  decline  during  The  time  a  are  nutrient  strong  or  available  autotrophic  source  That  discussed  bacteria  inorganic a  period  As  This  The  them  substances.  the  the  Phytoplankton before  of  energy  have  because  bacteria the  for  become  source  nutrient.  of  an  periods.  f i r s t  period.  group  phytoplankton  may  photosynthesis  bacteria  as  the  second  f i r s t  they  before  substance  the  component.  experiment  distinctive  biosynthesis.  compete  phytoplankton,  two  during  the  during  potentially  an  in  carbohydrate  nutrients  uptake.  growing  sections,  u t i l i z e  showed  are  u t i l i z e a l l  the  containing  As  the  experiment  restricted energy and  by  flow high  adaptive  as  results, range  there  bacteria in  I,  is  an  ecological  1 and  is  did  bacteria  1  gradient  salinity  gradient  separate  these  gradient.  If this  15a),  seawater freshwater gradient. this  However,  active  phytoplankton  grow.  a  heterogenous  e c o l o g i c a l components  temporal  distribution  of in  two  different  components  happens i n a body o f w a t e r moving  the  surface  l a y e r of the  estuary  spatial  distribution.  Numerous  heterotrophic  activity  low  range  1976;  salinity Wright,  concludes of  1978;  that  bacteria  in  there an  and  i t can studies  bacterial  o f an  be  expected  have  numbers  ( S t e v e n s o n and  et  1983).  al.,  i s a clear pattern estuary.  of  Estuaries  t o show a  indicated that  are  estuary  Wright  the  t o be  creates  two  the  middle of  end.  my  gradient  & Fig.  the  potential  of experiment before  In  Combining  from the  a  autotrophs,  salinity  salinity  many  energy.  ( F i g . 5a  the  rate  occupy  salinity  that  increased high  growth  f o r f r e s h and  survive  showed  period  salinity  may  inhibited.  is  pathways f o r  high  system  apparent  the  component  a v a i l a b l e to  II, part  not  mainly occupy  to temporally  the  not  internal  stage  i t  i n d i c a t e s t h a t the  along  bacteria  are  phytoplankton  first  condition  the  s u c c e s s f u l l y occupy the  component  heterotrophic  This  Because of  increasing salinity  but  during  have two  which are enough  autotrophic  system can  the  autotrophs  part  seawater  gradient  is  components,  autotrophic The  ability,  can  which  experiment which  some f a c t o r s , t h e  environments  long  i f the  simultineously.  different as  shows,  great  in  the  Erkenbrecher, Wright  spatial  (1983)  distribution  develop a  bacterial  100 flora  several  fresh  water  and in  or  Ferguson, the  salt  of  This  are  in  at  a result  these  the of  two  the  the  increase  show  between  i n d i c a t e d the  biological obvious.  stock a  of  correlation  only  standing  by stock  they  play  ecosystem. That  pathway  two  and  over  phenomena,  high  bacteria and  i n respones  among  and in  flux the  to  (Spies,  favours of  which  of  the  ecosystem  measuring we  energy flow  slow an  both  estimate  and  carbon  field  on  (Parker,  1982)  u n d e r what  down?  the  heterotrophic  (Parsons,  autotrophic  phytoplankton  to  i n the  1984)? A l s o ,  production  effects  under  r a t e can  s l o w down t o g i v e p r i o r i t y  generally  through  i s , u n d e r what c o n d i t i o n s d o e s  of n a t u r a l water  heterotrophic  flow  all  simultaneously  in big containers cultures  energy  i s a combination  1975),  inhibitory  the  to  b a c t e r i a , m i c r o f l a g e l l a t e s and  have i n d i c a t e d  the  transition  However,  workers  environment  specific  structure  the whole system  as  does  active  r e l a t e d t o each other  growth  laboratory  i n the  mouth  interrelations  role  many  (Palumbo  biologically  ecosystem  the  of  seems  Therefore  i n an  at  dynamics of  and  functional  to the  estuary  the  conditions.  can  compartments.  autotrophic  (1978)  head, a c t u a l l y a r e  standing  interactions  cycling  Wright  than e i t h e r  the  experiments. Therefore  pathways  phytoplankton  relations  of  b a c t e r i a are only  biomass  parallel  conditions  the  our  environmental The  The  is similiar  phytoplankton  population  concentration  s e a w a t e r a t e i t h e r end  estuary.  shown  high  in  bacteria declined significantly  water.  time  higher  1978).  upper  activity  the  fold  The  and  in  conditions inorganic  pathway  through  bacteria  (Lucas,  101 1955;  Sieburth,  Spies,  1984).  1959;  Considering biological  along  the  mouth  of  based  system,  seems  the  in  biomass  by  at  the  at  the  this  discharge marsh  head  intensity, bacterial  soil slow  a l l  inhibitors  nutrients,  response  advantage  an  to  be  this of  a l l along  fully  favourable  e.g.  by  r e l e a s e from  i . e . low  as  overcome  to  river  i n low  light  production  n u t r i e n t c o n c e n t r a t i o n as  growth  not  b a c t e r i a may  speed t o i n p u t of high  the  Many  resulting  o f b a c t e r i a such  and  at  level.  which are  sources,  low  behaviour  behaviour  indicate  turbidity  higher  of  with  t h e y may  the p o p u l a t i o n  of phytoplankton  nature  coefficient  on  possible  the p h y s i o l o g i c a l  a  phytoplankton  high organic n u t r i e n t supply  and  saturation  salinity  the  active  conditions  as  high  be  for  i n c r e a s i n g amounts  estuary,  show  etc.) higher growth  should  is a  considering  of estuary The  and  time  a h i g h l y productive area  least  b a c t e r i a are  (includes  base,  they at  space.  heterotrophic  In  However,  gradient  salinity  the  e t a l . , 1979;  residence  there  salinity  of  phytoplankton.  the  the  the ecosystem  head  component,  Kogure  a maximum a t h i g h  If  heterotrophic  utilize  low  estuary.  of  estuary,  reaching  unreasonable.  conditions  an  at  transition the  e t a l . , 1974;  limitation  production  phytoplankton  explored  Bell  high  of  well  uptake  rate  substances,  high  rate provide bacteria  the  time  limitation  by  circulation. In  a  real  components surface  estuary,  carried  layer  is  a  temporal  seaward always  in  a  accompanied  evolution moving by  an  of  water  ecological mass o f  the  increasing salinity  102 gradient.  For  level,  system  the  common, real  However, estuary  patterns,  experiment. current  effects,  substance  which  as  well  to  potential  activity  low s a l i n i t y Another that  as  the  happen  estuarine  sinking  that  i n our counter  phytoplankton  should  well  be h i g h e r  than  experiment  should  We s h o u l d  estuary  contrast,  at  the  expect  previous  attached  of  i n our simulated  salinity  and a r e a t t a c h e d  i n the last  found  decreased  reasonable  high  that a large organic  vigorousactivity  researchers  (1981)  bacteria a  Most  these  so t h e h e t e r o t r o p h i c  t h e b a c t e r i a numbers  p r o v i d e more  as I found  Albright  effects a l l  between o u r e x p e r i m e n t a n d f i e l d  known a n d i t i s r e a s o n a b l e  phytoplankton  bottom  release of  system  range.  our  of  and  a t t h e head o f an e s t u a r y  discrepancy  in  most  be  the  more  landward  trophodynamics.  them a r e b i g r o d s  may  outflow,  sedimentation  and  and  estuary  a n d marsh b a s e d c o a s t a l  degradation,  an  an  ina  circulation  may c o n t r i b u t e t h a n  bring  phytoplankton  bloom.  may be o v e r r i d d e n  In  sewage  may  estuary are i n  (e.g. d i f f e r e n t  etc.).  example,  processes  is  t h e m i c r o b i a l ecosystem  nature  factors  e n r i c h maximum t u r b i d i t y ,  contribute  is  on  a t t h e e s t u a r y mouth back t o t h e head o f t h e e s t u a r y  organic  at  common  many  organic  flow  and  i n o u r e x p e r i m e n t and i n r e a l  by  For  particles  process  this  tidal  allochthonous  to  this  end, a f t e r  increase greatly bacteria.  amount o f d e a d  substance  of attached  after the bacteria in  n o t found  this  zone.  i n the Fraser River estuary  phenomenon  This  p e r i o d o f our experiment.  have  along  studies  the s a l i n i t y in  a  real  In Bell  that the  gradient. estuary.  This In t h e  103 estuary, mouth,  when the  detritus  the  flow  will  water speed  sink  body slows  out  of  the  in  the  1985;).  The  b a c t e r i a attached  follow  free  the  living  bacteria can  in is  the  degradation  has  flasks. phase,  though  to  stop  sedimented (1984).  on  the  and is  a  sink.  the  attached  number o f  attached  range  the the  if  this  transportation  time f o r b a c t e r i a to compelete necessary  sink  find  a  of  great  if  transporting  after  stirrer,  great  nutrients  process.  out  that  showed  a  this  have t h i s  not  The  the the  attached  growth  i t was  amount o f d e a d  my  experiment with  separation  phytoplankotn  production  interaction  between along  i n both  the  experimental  tendency to  still  ths  process.  plateau  sediment. impossible  phytoplankton  T h i s a l s o happened i n S p i e s '  spatial  of  Thus  the  (LeBlond,  combining  result  l a y e r , but  estuary  the  bottom.  the  surface  will  head of  did  s i n k i n g and  Smetacek,  to  used a magnetic  Therefore  production  and  we  speed  component  the  can  flow  sinking  d i d not  the  circulation,  gradient  we  the  particulate material the  been c o m p l e t e d d u r i n g  phytoplankton we  of  phytoplankton  layer. This  increase  However  Even  the  not  salinity  population  the  on  the  surface  than the  experiments  dead  zone a t  e t a l . , 1983;  a c t u a l l y increases  low  shorter or  bacterial  the  This  degradation  our  of  The  (Lorenzen  would  frontal  u p p e r l a y e r as  s i n k i n g out  landward  comm.).  process  area  bacteria  carried  bacteria  In  host  s i n k out  be  pers.  plume  down.  decreases  also  reaches the  the  work  estuarine  heterotrophic the  salinity  ecosystemstructure  natural physical structure. In  estuaries,  a  sea  water  phytoplankton  based  ecosystem  104 mostly  occuppies  low  range  The  hetertrophic  sea  water  and  fresh  adaptation  for  the  patterns pool. are  of  the  are  At  salinity  released  as  inhibition  a  ecological  I  propose  physical gradient  may  be  bacteria.  made up  the  state  Their of  organic  the  of  the  amount  organic  the  phytoplankton  have  an  substance  substance,  of  both  evolutionary  physiological  of  the  bacteria  state  organic  of  substrate  bloom i n t e r m s o f  its  effect.  physical  structure  by  by  as  while  heterotrophic  spectrum.  c o n c e n t r a t i o n s of  well  gradient  These mixed p o p u l a t i o n s  whole s a l i n i t y  terms  Considering the  water.  affected in  by  (bacteria)  mainly decided  strongly  salinity  i s occuppied  population  limited  phytoplankton  high range of  a of space  (Fig.  the field space,  physical  and  the  and  conceptual estuarine by 25).  the  circulation  evolution  of  and  other  factors  in  e c o s y s t e m components  in  combining  t h e s e two  model  i n summary t o  ecosystem circulation  and  its  systems  together,  demonstrate  arrangement  p r o c e s s and  the  in  the the  conditional  Figure  25. The a r r a n g e m e n t i n an e s t u a r i n e  o f e c o s y s t e m components circulation field.  ^ o  106 Summary  1. in  In  the  an  upper  spatial  layer  certain  result  2.  gradient.  Most  organic  of  species  the  and  4.  internal  and  an  as  can  of the water  may  not  occur  i n an  the  pass through a salinity  may  mass.  estuary  not  in  function  as  but  salinity  increases,  occur  ecosystem  an  and  estuary.  autotrophic  they  contribute  q u i c k l y r e s p o n d and  grow i n  a maximum i n b i o m a s s a p p e a r s a t t h e gradient.  estuary  to  is  As  a  primarily occurs  result, the  a  high  autotrophic  result  of  a t some d i s t a n c e  marine  from  the  estuary. energy  flow  important energy  substances.  can  maximum p r o d u c t i o n  The  relatively  The  flagellates the  can  mortality  salinity  in  mouth o f an  return  die  estuarine  gradient  production  to  them  Seawater p h y t o p l a n k t o n  salinity  and  phytoplankton  an  movement  the  substances.  3.  end  in  the  ecosystem  seaward c o i n c i d e s w i t h  distributions  phytoplankton  components  d e v e l o p m e n t o f an  process.  mass  Freshwater  to  spatial  of  consequently  temporal  due  of t h i s  Freshwater  the  o f w a t e r moving  distribution  Therefore, as an  estuary,  carbon t o a low  if  which  t h r o u g h t h e m i c r o b i a l pathway the  can  microbial may  ecosystem be  created  has by  pathway w h i c h  have a g r e a t p o t e n t i a l  energy  flow  e n e r g y s t a t e as  and  allow  a  high  allochthonous i s made up ability  becomes level  organic  of b a c t e r i a  to contribute  the whole ecosystem  soon as p o s s i b l e .  of  to  107 5. on  A Salinity bacterial  gradient  growth. M i c r o f l a g e l l a t e s can a f f e c t  population  dynamics.  composition  within  to adapt 6.  substances then set  which  a  between  these  by  is  growth  the bacterial  their  species  a c l o s e s u c c e s s i o n and  mainly  microzooflagellates  process  decided  t o an u p p e r l i m i t ,  threshold.  two  shift  to obtain  component  allow  feeding  may  effect  conditions.  bacterial  exploited by  Bacteria  a population  to different The  does n o t seem t o have a s t r o n g  There  and  and t h e y a r e  a lower  i s no d y n a m i c  when o r g a n i c  by o r g a n i c  limit i s  equilibrium  energy sources a r e not  limited. 7.  A  hypothetical  proposed the  to  function  explain of  model  of  energy  level  field  theory i s  t h e dynamic s t r u c t u r e o f t h e e c o s y s t e m and  autotrophic  and h e t e r o t r o p h i c  components  i n an  ecosystem. 8. of well  Heterotrophic  the  estuary  production  and i s t h e r e s u l t  as t h e decay o f freshwater 9.  Heterotrophic  system,  being  bacterial  sinking  The  gradient  different  trophic  process.  over  but  is  more  be  expected  t h e mouth  substances as  i n the sea. ubiquitous  i n the  and m a r i n e s p e c i e s . at  the  a l s o a t c e r t a i n depths  area  A  of the  as a r e s u l t o f  materials.  estuarine  salinity  gradient  of allochthonous  phytoplankton  production  may  bloom  of organic  10.  maximally near  made up o f b o t h f r e s h w a t e r  maximum  phytoplankton  occurs  circulation  to  connect  transports the  materials  ecosystem  f u n c t i o n s which a r e separated  space.  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