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A field, culture, and cytological study of Porphyra gardneri, Porphyra nereocystis and Porphyra thuretii… Hawkes, Michael William 1978

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A F I E L D , CULTURE, AND CYTOLOGICAL STUDY OF POR PHYRA GARDNEHI, PQRPHYRA AND PORPHYRA  THURETII  (BHODOPHYTA,  IEREOCYSTIS  BANGIOPHYCIDAE)  by MICHAEL WILLIAM HAWKES .  Sc. (Honours), U n i v e r s i t y  of B r i t i s h  C o l u m b i a , 1973  A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE  REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY  in THE  FACULTY OF GRADUATE STUDIES (Botany  We a c c e p t to  THE  this  Department)  thesis  the required  as conforming standard  UNIVERSITY OF BRITISH COLUMBIA J u n e , 1978  ©  MICHAEL WILLIAM HAWKES, 197 8  In  presenting  this  an a d v a n c e d  degree  the  shall  I  Library  further  for  agree  scholarly  by  his  of  this  thesis at  the U n i v e r s i t y  make that  it  purposes  written  for  freely  permission may  representatives. thesis  in p a r t i a l  is  financial  of  of  Columbia,  British  available  for  for extensive  be g r a n t e d  It  fulfilment  by  shall  that  not  requirements I  agree  r e f e r e n c e and copying  t h e Head o f  understood  gain  the  of  be a l l o w e d  or  this  thesis  without  my  MICHAEL W. HAWKES  The  BOTANY  of  University  of  British  2075 Wesbrook Place Vancouver, Canada V6T 1W5  Date  9  MAY  1978  Columbia  or  publication  permission.  Department  tha  study.  my D e p a r t m e n t  copying  f<  ii  A biologist Humboldt  should s t r i v e  and D a r w i n , t o o b s e r v e  but above a l l , a t r i s k ever  t o be as d i v e r s e i n  wary o f dogma.  of b e i n g  with the i n t e n s i t y thought  an  outlook  as  of A g a s s i z ,  iconoclast,  to  be  iii  Research S u p e r v i s o r :  Dr.  Robert  F,  Scagel  ABSTRACT  A  field,  made o f  the  life  Bollenberg)  gpiphyra  number  basically  increased  host  spermatangium  and  P.  i s formed  layer  is  formed  In  P.  cells  by  February summer  autumn-winter  red  species.  of  undertaken  by  and  the  P.  carpogonium o f  to  The  and  i t s elongate,  and  reported  distribution  is  and  characterize  the  follow  their  P.  the can  of  a  In c o n t r a s t ,  nereocystis  spermatangia.  vegetative  gardneri  and  be  rectangular  probably  wall no  layer  new  wall  making  them  cells. P.  thuretii  bipolar  carpogonium only  division  and  new  cell.  because i t d i f f e r e n t i a t e s  nereocystis  species,  spermatogenesis to  production  vegetative  by  through  species  geographical  study  to d i s t i n g u i s h from  prctctrichogynes  thuretii  epiphytic  spermatangium i n Porphyra g a r d n e r i  a transformed  recognized  from  spring  carpogonium  The  difficult  similar  P.  et  species.  was  around  (Smith  and  spring-early  and  microscope  seguence. thuretii  a an  species  carposporogenesis  The  is  f o r a l l three  light  gardneri  Anderson  found  i s a winter-early  of  Porphyra  superficially  was  nereocystis  thuretii  A  is  of  been  Bangiales).  gardneri  but  c y t o l o g i c a l i n v e s t i g a t i o n has  nereocystis  Dawson, t h r e e  Porphyra  £•  P.  (Bhodophyceae,  November,  and  histories  Hawkes,  Setchell et algae  culture  easily  prototrichogynes.  does  d i s t i n g u i s h e d from shape.,  is  not  form  vegetative  iv  On  the  basis of these  literature separated and  I  suggest  carpogonia  g r o u p s based the  first  carposporogenesis respect  t o the  The  is  in  both  Of  the  produces  formation  three  and  species  monospores.  an  abundant  small  is  level  large  observations  photoperiod formation  is  an  important  o f s p e r m a t a n g i a and  A conchocelis  the  by  the  vesicles. are  In  P.  observed  division.  and  gardneri  temperature  foliose  phase.  production The  in of  economic  suqqest  that  factor involved i n inducinq  the  carpogonia. for  the  Porphyra t h u r e t i i .  temperature reqimes t e s t e d carpospores  conchocelis  Hollenberq  discussed.  Porphyra g a r d n e r i  phase i s r e p o r t e d  RQ.I2hlJL<* 2 § £ ^ a S £ i and and  of  et  monosporogenesis  fibrous  i m p l i c a t i o n s o f monospore p r o d u c t i o n Field  into  with  workers.  Porphyra  photoperiod  characterized  and  other  Smith  only  and  discussed  carposporangial  a l l  ultrastructural  Porjphjfra g a r d n e r i  is  on t h e b a s i s o f t h e  r e g i m e s t e s t e d monospores g e r m i n a t e d At  defined  w a l l l a y e r , whereas  This  studied,  Under  has  spermatogenesis  t r a n s f e r of Porphyrella Gardneri  mode o f c a r p o g o n i u m  group  clearly  s e q u e n c e s r e p o r t e d by  Porphyra g a r d n e r i i s explained  be  features.  periclinal.  division  the  spermatangial  advanced and  in  Porphyra can  their  d e p o s i t i o n of a new  division  reports  of  on  more  prototrichogynes  t o the  and  species  more p r i m i t i v e group l a c k s t h e s e The  to  two  the  morphology;  with  s p e r m a t a n g i a due the  that  into at least  carpogonial  observations  first  time  for  Under a l l p h o t o p e r i o d qerminated  into  the  phase. sardneri  and  P.  thuretii  short  days  favoured  V  ccnchosporanqial photoperiodic my  results  indicate  branch  effect were  that  was  formation;  observed  f o r P.  inconclusive,  temperature  however, nereocystis.,  reports  i s a critical  no  in  the  factor  such  Although  literature  i n ccnchospore  release. Amoeboid  movement  E°I£J3J£§ t h u r e t i i of  this  of  and d i s c u s s e d  P.  basal of  nereocystis rhizoids  was  examined  blade,  The s p e r m a t i a examined  ultrastructurally  Florideophycidae  Porphyra g a r d n e r i microscopes.  occurrence  them  nereocystis  Porphyra  gardneri  microscope.  deep i n t o P.  The  the medulla nerepcystjs  of the Nereocvstis  g a r d n e r i and P.  stipe.  nereocystis  and f o u n d t o have many  and v e g e t a t i v e  in  with  cells  T h i s i s the f i r s t  were  similarities  the  life  history  the l i g h t  and e l e c t r o n  and also  in light  attached  report of  t o the carpogonia  canals  P. t h u r e t i i occurs  i n their  i n other of my  of the conchocelis  unequivocal  of  for phase  sexual  Porphyra.  fertilization  reproduction  evaluated  penetrate  documented  i n t h e genus  via  reproduction  of  spermatia.  Suspected spermatia  sexual  for  A d i p l o i d chromosome number 2n=8 i s r e p o r t e d  °f £• g a r d n e r i .  P.*  to the  the l i g h t  of f e r t i l i z a t i o n is  carpospores  reproduction  with  penetration  of jgorphyra  The o c c u r r e n c e  to  reported  whereas t h e r h i z o i d s o f  make o n l y a s u p e r f i c i a l  the  respect  attachment zone  o f P. g a r d n e r i  the L a m i n a r i a  with  with  is  phenomenon i n t h e R h o d o p h y t a g e n e r a l l y .  The e p i p h y t e - * h o s t * and  conchospores  have  been  sugqestinq life  and  linked  observed that  histories.  sexual  Reports of  members o f t h e B a n q i o p h y c i d a e  observations.  in  are  vi  TABLE OF CONTENTS  AESTRACT  i i i  TABLE OF CONTENTS L I S T OF TABLES  vi  ••••••••*•«••••••••* * * ••*•*«***•••••«*••**«*  LIST OF FIGURES  XX x i i XXV  L I S T OF APPENDICES ACKNOWLEDGEMENTS  ...  . xxvi  GENERAL INTRODUCTION  1  TERMINOLOGY  4  ......................... . . ....................  PART I - FIELD  AND HERBARIUM STUDIES  ......................  9  ...........................................  9  M a t e r i a l s a n d methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  14  Results  17  Introduction  ................................................  *• R.Q££fel£.§ g a r d n e r i  ...................................  1. S e a s o n a l o c c u r r e n c e 2.  *Host*  species  3. A r t i f i c i a l  host  o f e p i p h y t e and ' H o s t *  .......  17  ...................................  20  experiment  .......................  21  ........................  21  ................................  23  4. G e o g r a p h i c a l d i s t r i b u t i o n B» P o r p h y r a n e r e o c y s t i s  1. S e a s o n a l o c c u r r e n c e o f e p i p h y t e and ' H o s t 2. V e r t i c a l 3.  'Host  1  distribution  .......  23  ............................  26  1  species  4. A r t i f i c i a l  host  17  26 .......................  26  ........................  27  t h u r e t i i ...................................  28  1. S e a s o n a l o c c u r r e n c e . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  28  5. G e o g r a p h i c a l C. P o r p h y r a  experiment  distribution  2. H o s t * s p e c i e s ... 1  3. A r t i f i c i a l  •Host*  experiment  4. G e o g r a p h i c a l d i s t r i b u t i c n Discussion  ..........  ............,  .... .... ... . .  *• P o r p h y r a  g a r d n e r i .....  E, P o r p h y r a  nereocystis.,  C. P o r p h y r a  t h u r e t i i ,,.,. AND C A EPOSPOROGEN ESIS  FART I I - SPERMATOGENESIS Introduction  .............  M a t e r i a l s and methods .... S€SUltS  *•* *•••a*** #••* •• •  A. P o r p h y r a  g a r d n e r i .....  •• •  1. S p e r r a a t o g e n e s i s .•.. 2. C a r p o s p o r o g e n e s i s .. 3. E x a m i n a t i o n o f t y p e l o c a l i t y B. P o r p h y r a  n e r e o c y s t i s ..  1. S p e r m a t o g e n e s i s  ....  2. C a r p o s p o r o g e n e s i s .. C. P o r p h y r a 1.  t h u r e t i i .....  Spermatogenesis  ....  2. C a r p o s p o r o g e n e s i s .. Discussion  .. • • • . . . . . . . . . .  PABT I I I - CULTURE STUDIES Introduction  . . . . . . . . • •...  M a t e r i a l s and methods . . . . Results  ... * . . . *..•"•.. *..  A. P o r p h y r a 1.  g a r d n e r i .....  Monospores  .........  p l a n t s ...  viii  2. Spermatangium  and c a r p o g c n i u w  formation  3. C a r p o s p o r e g e r m i n a t i o n - t h e c o n c h o c e l i s 4. C o n c h c s p o r a n g i a l  branch  ..........  95  phase ...  95  formation  97  5. C o n c h o s p o r e r e l e a s e B  « <i!PJE£lLY££ n e r e o c y s t i s  97  ................................  1. C a r p o s p o r e g e r m i n a t i o n 2. C c n c h o s p o r a n g i a l  C  - the c o n c h o c e l i s  branch formation  phase  98  ...  98  ................  100  3. C o n c h o s p o r e r e l e a s e  100  RQI2k2L& llJM£gtii . . . . . . v . . . . . . . . . . . . . . . . . . . . . . . . . . . .  101  1. C a r p o s p o r e g e r m i n a t i o n  phase ...  101  ................  103  ..............................  103  .............................................  104  - the conchocelis  2. C c n c h o s p o r a n g i a l b r a n c h f o r m a t i o n 3. C o n c h o s p o r e r e l e a s e Discussion  1. Monospores 2. Spermatangium 3.  Effect  and c a r p o g o n i u m f o r m a t i o n  of photoperiod  germination  on monospore and  .  104  ..........  105  carpospore  •.....................................  4..The c o n c h o c e l i s  phase  ............................  106 107  5. C o n c h o s p o r a n g i a l b r a n c h e s  111  6.  113  Conchospore r e l e a s e  .*  7. C c n c h o s p o r e s ....... . . . . . • . . . . . . . . . . . . . . . . . . . . . . . . PAST IV - CYTCLCGICAL STUDIES General introduction  116 128  ..... . . . . • . . . . . . . . . . . . . . . . . . . . . . . . .  128  M a t e r i a l s and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  128  A. E p i p h y t e - * H o s t ' a t t a c h m e n t  ..........................  131  ...........................................  131  Introduction Besults 1*  .......  Porphyra g a r d n e r i  ..  132  . . . . . . . . . . . . . . ..... .............  132  ix  2. P o r p h y r a  . . . . . . . . . . . « . . . ..............  132  ....................... ......................  133  B . M o n o s p o r o g e n e s i s ........... .. ... ... ... . . . . . . . . . . ... •  134  Introduction  134  Discussion  BesuIts  ..............».«•»....»»» . . . . . . . . . . . . . . . . . .  . '...".*...... . ....."'..« •• * ...... .. •......... .<**.'.«*•*  Discussion C.  nereocystis  .....•»• .• '•*...... . .... . .... . . ...... . ... . . ...... . . 136  Sexual reproduction  Introduction  ............  137  ...........................................  137  1* The spermatiutn 2.  134  i n Porphyra gardneri  ...................................  139  Reports o f f e r t i l i z a t i o n .........................  141  3. R e p o r t s o f f u n g a l i n f e c t i o n Results  of Porphyra  ..........  143  . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ...  146  1. S p e r m a t o g e n e s i s - e l e c t r o n m i c r o s c o p y microscopy  ............  146  2. F e r t i l i z a t i o n  - light  .................  147  3. F e r t i l i z a t i o n  - e l e c t r o n microscopy ..............  148  4. Chromosome c o u n t s 5.  Feulgen s t a i n i n g  Discussion  150 . . . . . * . . . . * . . . . . . . . * . ..**....•...  * •••••>'•'•"» »•'..". »'«»«»«>».•••»•». «•'...•* .......  ...•« •••..  150 152  1. S p e r m a t o g e n e s i s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  152  2. F e r t i l i z a t i o n  ...••  155  ... .......  160  .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ......  160  . . . • • . . . . •. • •..... .....  D. E v i d e n c e o f s e x u a l  reproduction  Porphyra n e r e o c y s t i s Introduction Results  . . . v.'.  and P o r p h y r a t h u r e t i i  . ... «•.... .  1. P o r p h y r a n e r e o c y s t i s 2.  GEN ERA!  ' . - . •'.  . . . . . . . . . . . .• «•.'••....  160  ...................... .......  160  . . . . . . . . . . . . . . . . . . . . . . ..........  161  . . . . . . . . . . . .... . . . . . . . . . . . . . . . . . . .. ........ . .  162  Porphyra  Discussion  in  SUMMARY  ..  thuretii  . . . . . . . .... . . . . . . . . . ... .V . . . . . . . . . . . . . . . . ,  209  X II11BATDEE CITED  212  APPENDIX I ........... . . . . . . . . . . . . . . . . . . . . . . . . . .......... ..  234  APPENDIX I I . . . . . . . . . . . . . . . . . . . . . . . . . ........... ...........  236  xi  LIST  I. II. III.  IV, V.  VI.  VII.  Herbaria  f r o m which  OF  TABLES  specimens  were e x a m i n e d  ......  Host p l a n t s o f P o r p h y r a g a r d n e r i Biomass (grams d r y w e i g h t ) o f P o r p h y r a n e r e o c y s t i s per v e r t i c a l Nereocystis  s t i pe  Host p l a n t s  of Porphyra t h u r e t i i  39  meter  of  ............................... • .................  Annual v a r i a t i o n i n d u r a t i o n o f d a y l i g h t {hr. and min.) on t h e 2 1 s t day o f t h e month a t 35°N and 48° 50.1«N ( s t u d y s i t e ) . . . . . . . . . . . . . . . . . Summary o f t h e s p e c i e s o f P o r p h y r a w h i c h p o s s e s s p r o t o t r i c h o g y n e s , and t h o s e w h i c h do not  40 41  42  74  A n n u a l t r e n d i n s u r f a c e w a t e r t e m p e r a t u r e (°C) a t t h e s t u d y s i t e (48° 50.1»N, 1 2 5 ° 11.1»W) and A m p h i t r i t e P o i n t (48° 5 5 . 2 * N , 125° 32.2 W) ...  119  The s p e c i e s o f P o r p h y r a f o r w h i c h d i p l o i d chromosome numbers have been r e p o r t e d f o r t h e c a r p o s p o r e s o r v e g e t a t i v e c e l l s of the c o n c h o c e l i s phase  163  f  VIII.  38  xii  L I S T OF FIGORES  1a.  Monosporic t h a l l i c f Porphyra g a r d n e r i growing on t h e b l a d e m a r g i n s o f L a m i n a r i a s e t c h e l l i i . The t y p e s p e c i m e n (DS 306401 i n UC) . . . . . . . . . . . . . . .  44  A 2.7 m# f e r t i l e s p e c i m e n o f Porphyra n e r e o c y s t i s c o l l e c t e d a t the study s i t e (UBC 57180) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  44  1c.  The l e c t o t y p e  44  2a.  The f i e l d work was c a r r i e d o u t i n B a r k l e y Sound on t h e west c o a s t o f V a n c o u v e r I s l a n d ( i n s e t ) . The s t u d y s i t e was l o c a t e d o f f D i a n a I s l a n d ( c i r c l e d ) , near t h e B a m f i e l d Marine S t a t i o n (from C a n a d i a n H y d r o g r a p h i c S e r v i c e C h a r t #L-3001) ...  46  A e r i a l p h o t o g r a p h o f D i a n a I s l a n d and t h e s t u d y s i t e . The d i r e c t i o n o f t h e open P a c i f i c Ocean i s i n d i c a t e d by t h e a r r o w , (from B. C. .Government a i r p h o t o g r a p h #BC 7238-182) .....  46  C h a r a c t e r i s t i c h e a r t - s h a p e d new b l a d e g r o w t h o f L a m i n a r i a s e t c h e l l i i (UBC 54895) . . . . . . . . . . . . . . . . . .  48  E a r l y s t a g e i n t h e s p l i t t i n g of the b l a d e o f L. s e t c h e l l i i (UBC 57204) . . . . . . . . . . . . . . . . . . . . . . . . .  48  Mature, l a c e r a t e d b l a d e o f L. s e t c h e l l i i with t h a l l i of P o r p h y r a g a r d n e r i a l o n g i t s margins (UBC~54896) . . . . . . . . . . . . . . . . . . . . . . . . . . .  48  Monosporic t h a l l i  48  1b.  2b.  3a. 3b. 3c.  3d.  o f Porphjrra t j u £ § t i i  o f P. jgarjner_i Porphyra  4a.  4b.  4c-e.  (UC 791973)  {UBC  54819)  ....  ......  gardneri  T h a l l u s o f P o r p h y r a g a r d n e r i which h a s s p e r m a t a n g i a , c a r p o g o n i a and young carposporangia a l o n g i t s l a t e r a l margins. Monospores a r e s t i l l b e i n g r e l e a s e d a l o n g t h e d i s t a l margin (UBC 54821)  50  O l d , l a c e r a t e d , and e p i p h y t i z e d t h a l l u s o f L a m i n a r i a s e t c h e l l i i with a few t h a l l i o f P o r p h y r a g a r d n e r i on i t s m a r g i n s ( a r r o w s ) . The new L a m i n a r i a b l a d e i s j u s t b e g i n n i n g t o grow (UBC 57205) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  50  Morphological v a r i a t i o n i n t h a l l i of P. g a r d n e r i w h i c h a r e s p e r m a t a n g i a l and c a r p o s p o r a n g i a l . Note t h e c h a r a c t e r i s t i c p a t t e r n f o r m e d by t h e s p e r m a t a n g i a a l o n g t h e  xiii  4f.  l a t e r a l m a r g i n s , f i g u r e 4 c : UBC 54138, f r o m B o t a n i c a l Beach; f i g u r e s 4d ana 4es UBC 54891, f r o m Cape B e a l e . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  50  T h a l l u s o f Pi g a i r d n e r i f r o m t h e t y p e l o c a l i t y . T h i s p l a n t i s r e l e a s i n g monospores a l o n g i t s d i s t a l m a r g i n a n a s p e r m a t i a ana c a r p o s p o r e s a l o n g i t s l a t e r a l m a r g i n s (UBC 54818) . . . . . . . . . . . . .  50  Porphyra 5a.  5b.  5c.  5d.  5e.  nereocystis  Young v e g e t a t i v e t h a l l i a r e c h a r a c t e r i s t i c a l l y l o n g ana s t r a p s h a p e d , t a p e r i n g g r a a u a l l y t o a point (DBC 57187)  52  The f o r m a t i o n o f s p e r m a t a n g i a b e g i n s a t t h e l a t e r a l m a r g i n s and p r o c e e d s b a s a l l y . These r e g i o n s o f t h e t h a l l u s a r e e a s i l y r e c o g n i z e d by t h e i r p a l e y e l l o w t o w h i t e c o l o u r (OBC 57188) .....  52  as a r e s u l t o f s p e r m a t i u m l i b e r a t i o n , t h e l a t e r a l s p e r m a t a n g i a l margins begin t o erode. T h i s s t a r t s a t t h e apex and p r o c e e d s b a s a l l y . The r e m a i n i n g a p i c a l p o r t i o n o f t h e t h a l l u s c o n s i s t s o f v e g e t a t i v e c e l l s and c a r p o s p o r a n g i a (UBC 57184)  52  S p e r m a t a n g i a f o r m i n g m a r g i n a l and s u b m a r g i n a l s t r e a k s w h i c h a r e o r i e n t e d i n an apex t o base d i r e c t i o n (UBC 57180) . . . . . . .  52  The t i p s o f o l d e r t h a l l i o f t e n have a s l i g h t l y m o t t l e d a p p e a r a n c e c a u s e d by abundant c a r p o s p o r a n g i a (UBC 57192) . . . . . . . . . . . . . . . . . . . . . . . .  52  Porphyra 6a.  6b.  6c,d.  6e.  thuretii  M o r p h o l o g i c a l v a r i a t i o n i n t h a l l i which were g r o w i n g on t h e s t i p e o f N e r e o c y s t i s l u e t k e a n a i n B a r k l e y Sound. Note t h e c h a r a c t e r i s t i c r u f f l e d m a r g i n s (UBC 57200) . . . . . . . . . . . . . . . . . . . . . . .  54  The p r o d u c t i o n o f s p e r m a t a n g i a b e g i n s i n t h e a p i c a l marginal r e g i o n of the t h a l l u s (UBC 52109) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  54  On o l d e r t h a l l i s h o r t s t r e a k s o f s p e r m a t a n g i a f o r m s u b m a r g i n a l l y . Note t h e c h a r a c t e r i s t i c p a t t e r n t h a t r e s u l t s (UBC 57201 and UC 95596 respectively)  54  Specimen from t h e M o n t e r e y P e n i n s u l a which r e s e m b l e s s p e c i m e n s f r o m B a r k l e y Sound i n s i z e and s p e r m a t a n g i a l p a t t e r n •  54  xiv  Porphyra 7a.  7b.  7c.  gardneri - spermatogenesis, l i g h t microscopy  V e g e t a t i v e c e l l w i t h l a r g e c e n t r a l p y r e n o i d and c h l o r o p l a s t . The n u c l e u s i s p e r i p h e r a l l y l o c a t e d .;,........................................  77  New w a l l l a y e r (arrow) l a i d down a r o u n d spermatangium p r i c r t o the f i r s t d i v i s i o n .........  77  The f i r s t periclinal  7d-f. 7g«  d i v i s i o n o f t h e spermatangium i s ........................................  S u b s e g u e n t d i v i s i c n s of t h e s p e r m a t a n g i u m Mature spermatangium  .........  7h.  S u r f a c e view o f an immature s p e r m a t a n g i u m  7i.  Surface  8a.  8b.  77  4 s p e r m a t i a deep i n  ^ jr&nsv*sxT3*3 ssc"bxon • • • * * •* • • *» • • * * • • * • • • • • * • * • * • * • « •  view  77  of a mature  spermatangium  .........  77  ............  77  Porphyra g a r d n e r i S p e r m a t o g e n e s i s . The d i v i s i o n s e q u e n c e l e a d i n g t o a mature s p e r m a t a n g i u m o f 64 s p e r m a t i a . Note t h e p r o d u c t i o n o f a new w a l l l a y e r ( i n d i c a t e d by a t h i c k e r l i n e ) p r i o r t o d i v i s i o n , a n d t h e absence o f any • c r u c i a t e * d i v i s i o n s ............... C a r p o s p o r o g e n e s i s . D i v i s i o n s e g u e n c e which p r o d u c e d 2, 4 o r 8 c a r p o s p o r e s per c a r p o s p o r a n g i u m . The u p p e r c a r p o s p o r a n g i u m c o n t a i n i n g 4 c a r p o s p o r e s i s most o b v i o u s i n s u r f a c e view and i s shown i n F i g u r e s 9g and 9h. The d i v i s i o n pathway i n d i c a t e d by s o l i d a r r o w s i s most common . , , . . « ^ .......'........*........ .< Porphyra j a r d n e r i -  *7 *7  79  79  carposporogenesis  9a.  Surface  ..................  81  9b.  S u r f a c e v i e w o f s p e r m a t a n g i a (lower l e f t ) , c a r p o g o n i a and c a r p o s p o r a n g i a . . . . . . . . . . . . . . . . . . . . .  81  T r a n s v e r s e s e c t i o n o f t h e t h a l l u s s h o w i n g two c a r p o g o n i a . Note t h e b i p o l a r p r o t o t r i c h o g y n e s and a t t a c h e d s p e r m a t i u m (arrow) . . . . . . . . . . . . . . . . . . .  81  T r a n s v e r s e s e c t i o n s h o w i n g two c l o s e l y a p p r e s s e d c a r p o g o n i a . Note a t t a c h e d s p e r m a t i u m (arrow) . . . . . . . . . . . . . . . . . . . . . •..... . . . . . . . . . . . .....  81  T r a n s v e r s e s e c t i o n showing t h a t t h e f i r s t d i v i s i o n o f t h e caposporangium i s p e r i c l i n a l  81  9c.  9d.  9e, 9f.  view o f v e g e t a t i v e c e l l s  Transverse  section  showing  the o r i e n t a t i o n  of  ......  XV  t h e t h i r d and f o u r t h d i v i s i o n s of the c a r p o s p o r a n g i u m . Shown i n s u r f a c e v i e s i n F i g u r e s 9g a n d 9h. See a l s o F i g u r e 8a . . . . . . . . . . . . .  81  S u r f a c e view o f c a r p o s p o r a n g i u m t h a t has u n d e r g o n e t h r e e d i v i s i o n s and c o n t a i n s 4 c a r p o s p o r e s . See a l s o F i g u r e 8a . . . . . . . . . . . . . . . . . . .  81  9h.  As i n F i g u r e  ...  81  9i.  Transverse s e c t i o n through a carposporangium c o n t a i n i n g 8 c a r p o s p o r e s ( o n l y 4 v i s i b l e ) .........  81  S u r f a c e view o f a c a r p o s p o r a n g i u m c o n t a i n i n g 8 c a r p o s p o r e s ( o n l y t h e f o p 4 a r e v i s i b l e ) ..........  81  9g.  9j.  9g, b u t a d i f f e r e n t  Porphyra 9k.  91.  S u r f a c e view  c f s p e r m a t a n g i a which  a few d i v i s i o n s  Porphyra  10b.  10c.  1.0e.  10f.  10g. 10h.  .........................  81  nereocystis - spermatogenesis  Transverse section  Porphyra  81  have  T r a n s v e r s e s e c t i o n t h r o u g h an immature spermatangium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . , S u r f a c e view of mature s p e r m a t a n g i a l r e g i o n o f the t h a l l u s . I n d i v i d u a l spermatangia are not c l e a r l y d e l i m i t e d .................................  spermatangium  10d.  nereocystis - spermatogenesis  S u r f a c e view o f t h e f o l i o s e p h a s e s h o w i n g v e g e t a t i v e c e l l s with a s i n g l e s t e l l a t e c h l o r o p l a s t .......................................  undergone  10a.  plane o f f o c u s  83  t h r o u g h a mature  which i s 8 s p e r m a t i a nereocystis  deep  ...........  83  - carposporogenesis  Transverse s e c t i o n through a vegetative c e l l r e g i o n o f t h e t h a l l u s . Note t h e s i n g l e s t e l l a t e c h l o r o p l a s t , c e n t r a l p y r e n o i d and l a t e r a l n u c l e u s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .............. T r a n s v e r s e s e c t i o n t h r o u g h a c a r p o g d n i u m . Note i t s s l i g h t l y more e l o n g a t e shape compared t o the v e g e t a t i v e c e l l s T r a n s v e r s e s e c t i o n s h o w i n g two c l o s e l y a p p r e s s e d c a r p o g o n i a . Note t h e spermatium (arrow) and f e r t i l i z a t i o n c a n a l T r a n s v e r s e s e c t i o n o f two c a r p o s p o r a n g i a shewing t h a t the f i r s t d i v i s i o n i s p e r i c l i n a l Transverse section  83  o f two c a r p o s p o r a n g i a . The  83 83  83 .....  83  xvi  10i.  10j.  10k.  101.  11a.  one on t h e l e f t i s a t t h e s t a g e shown i n s u r f a c e view i n F i g u r e 1 0 i , whereas t h e one on the r i g h t i s a t t h e s t a g e shown i n F i g u r e 1 0 j . T h i s d i v i s i o n s e g u e n c e i s t h e same a s t h a t shown f o r P o r p h y r a g a r d n e r i i n F i g u r e 8b  83  S u r f a c e view o f a c a r p o s p o r a n g i u m c o n t a i n i n g 4 c a r p o s p o r e s .......................................  83  S u r f a c e view of a c a r p o s p o r a n g i u m c o n t a i n i n g ( o n l y t h e t o p 4 a r e v i s i b l e ) . See a l s o F i g u r e 10h  83  8  S u r f a c e view o f mature c a r p o s p o r a n g i a . I t i s d i f f i c u l t t o determine the boundaries of a s i n g l e carposporangium ............................  83  T r a n s v e r s e s e c t i o n s h o w i n g two c l o s e l y a p p r e s s e d c a r p o s p o r a n g i a which have u n d e r g o n e more t h a n one d i v i s i o n i n t h e p l a n e o f t h e t h a l l u s ...........................................  83  Porphyra n e r e o c y s t i s , t r a n s v e r s e s e c t i o n t h r o u g h a r e g i o n o f mature c a r p o s p o r a n g i a . I t i s d i f f i c u l t t o determine the o r i g i n a l b o u n d a r i e s o f a s i n g l e carposporangium ............  85  Porphyra  t h u r e t i i •• - s p e r m a t o g e n e s i s  11b,c.  11d. 11e.  11f.  S u r f a c e view showing e a r l y d i v i s i o n s o f t h e s p e r m a t a n g i a . . . . . . . . . . . . . . . . . . . . . . ...............,  85  T r a n s v e r s e s e c t i o n t h r o u g h an immature spermatangium .....................................  85  Mature s p e r m a t a n g i u m w i t h s u r f a c e d i m e n s i o n s a/4, b/4 and c o n t a i n i n g a t o t a l o f 64 s p e r m a t i a . ........................................  85  T r a n s v e r s e s e c t i o n t h r o u g h a mature s p e r m a t a n g i u m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . , .,  85  Porphyra t h u r e t i i 11g.  11h. 11i.  -  carposporogenesis  T r a n s v e r s e s e c t i o n t h r o u g h a carpogonium which a p p e a r s t o have v e r y s l i g h t b i p o l a r p r o t o t r i c h o g y n e s . . . . . . . . . . . . . . . . . ....... T r a n s v e r s e s e c t i o n showing t h a t t h e f i r s t d i v i s i o n of t h e carposporangium i s p e r i c l i n a l  85 .....  Transverse s e c t i o n o f a carposporangium t h a t has undergone s e c o n d and t h i r d a n t i c l i n a l d i v i s i o n s . A s u r f a c e view i s shown i n F i g u r e 11j ........................................  85  85  xvii  11j. 11k.  111.  S u r f a c e view o f a c a r p o s p o r a n g i u m w h i c h undergone t h r e e d i v i s i o n s  35  S u r f a c e v i e w o f a mature c a r p o s p o r a n g i u m c o n t a i n i n g 8 carpospores (only the top 4 a r e v i s i b l e ) . See f i g u r e 111 . . . . . . . . . . . . . . . . . . . . . . . . . .  85  Transverse section  85  o f a m a t u r e c a r p o s p o r a n g i u m ....  Porphyra 12a.  has  gardneri  Mcnospores b e i n g r e l e a s e d a l o n g the margin o f a t h a l l u s c o l l e c t e d a t t h e study s i t e ...............  121  V a r i o u s s t a g e s i n the development of the f o l i o s e t h a l l u s from a monospore. N o t e t h e l o n g r h i z o i d a 1 p r o t u b e r a n c e s ...........................  121  12g.  Released  121  12h,  Carpospore germination to give the conchocelis phase . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  12b-f.  12i.  Single spore-like swelling filament  12j.  carpospores  branches ......................... Por£h_y.ra  13a. 13b. 13c.  on t h e c o n c h o c e l i s  ............................................  Conchosporangial  121  121 121  gardneri  C o n c h o c e l i s f i l a m e n t s , c o n c h o s p o r a n g i a l branch ( l o w e r r i g h t ) and r e l e a s e d c o n c h o s p o r e (arrow) .... B i p o l a r s p o r e l i n g r e s u l t i n g from conchospore germination ....  123  C h a r a c t e r i s t i c p i n n a t e l y b r a n c h e d morphology o f t h e c o n c h o c e l i s phase g r o w i n g i n o y s t e r s h e l l .....  123  Porphyra  nereocystis  13d.  Beleased  ..............................  123  13e.  Carpospore g e r m i n a t i o n to g i v e the c o n c h o c e l i s phase .............................................  123  Basal s p o r e - l i k e swelling of the conchocelis phase .............................................  123  13f. 13g.  Apical  carpospores  123  spore-like  swelling  of the c o n c h o c e l i s  phase .............................................  123  13h.  I r r e g u l a r and  beaded-branch t y p e s  .................  123  13i.  Sguare-celled  branch ..............................  123  xviii  13 j .  Small ccnchosporangial vegetative filament  branch connected t o the  by a s g u a r e - c e l l e d  b r a n c h .....  13k.  Conchosporangial  branches  14a.  Porphyra n e r e o c y s t i s C l o s e u p o f c o n c h o s p o r a n g i a l b r a n c h e s showing t h e s i n g l e s t e l l a t e c h l o r c p l a s t i n each ccnchosporangium ................................  123 123  125  14b.  P i t p l u g s ( a r r o w s ) between c o n c b o s p o r a n g i a i n the c c n c h o s p o r a n g i a l branch ( t r e a t e d with chromosome f i x a t i v e and s t a i n ) . . . . . . . . . . . . . . . . . . . . . 125  14c.  Divisions within  b r a n c h ......  125  14d.  C h a r a c t e r i s t i c pinnate branching of the c o n c h o c e l i s p h a s e g r o w i n g i n o y s t e r s h e l l . Note the i r r e g u l a r s w e l l i n g s ...........................  125  the c o n c h o s p o r a n g i a l  Porphyra 14e.  Released  ..............................  125  14f.  Carpospore g e r m i n a t i o n to g i v e the c o n c h o c e l i s phase .............................................  125  V e g e t a t i v e c o n c h o c e l i s f i l a m e n t s , beaded b r a n c h e s and young c o n c h o s p o r a n g i a l b r a n c h (arrow) . . . . . . . . . . . . . . . . . . . . . . . .... . . . . . . . . . . . . . . . .  125  C l u s t e r o f c o n c h o s p o r a n g i a l b r a n c h e s showing s i n g l e s t e l l a t e c h l o r o p l a s t i n each conchosporangium ..................................  125  14g.  14h.  14i.  carpospores  thuretii  Candelabra-shaped c l u s t e r branches showing t h e i r  of c o n c h o s p o r a n g i a l  positive  phototropism  ......  125  14j.  C c n c h o s p o r a n g i a l b r a n c h w i t h c o n c h o s p o r e s .........  125  14k.  Released  125  conchospores ............................. Porphyra t h u r e t i i  15a-d.  15e.  C o n c h o s p o r e s e x h i b i t i n g amoeboid change o f s h a p e and movement. F i g u r e s 15a and 15b t a k e n 90 s e c o n d s a p a r t . F i g u r e s 15c and 15d t a k e n 45 seconds apart ......................................  127  Young b i p o l a r s p o r e l i n g r e s u l t i n g from c o n c h o s p o r e g e r m i n a t i o n . Note r h i z o i d a l protuberance ......................................  127  xix  151,g.  15h. 15i, j .  16a.  O l d e r , u n i s e r i a t e s p o r e l i n g s s t a r t i n g t o form b a s a l pad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  127  Biseriate  127  stage  of s p o r e l i n g  growth  O l d e r s p o r e l i n g s . Note t h e b a s a l r h i z o i d a l pad i n F i g u r e 15 j . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  127  T r a n s v e r s e s e c t i o n o f the blade margin o f L a m i n a r j a s e t c h e l l i i and t h e b a s e o f P o r p h y r a g a r d n e r i . The r h i z o i d a l f i l a m e n t s c a n be s e e n p e n e t r a t i n g i n t o t h e m e d u l l a o f t h e L a m i u a r i a blade ..............................  166  16 fe. C l o s e u p o f t h e r h i z o i d a l p e n e t r a t i o n i n t o t h e m e d u l l a . . . . . . . . . . . . . . ... .......... .......... 16c. 16d. 17.  18.  19a. 19b.  19c.  166  Transverse s e c t i o n through a Nereocystis s t i p e and t h e P o r p h y r a n e r e o c y s t i s a t t a c h e d t o i t .......  166  C l o s e u p of the c e n t r a l r e g i o n o f the attachment d i s c . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . .  166  Tranverse s e c t i o n of Porphyra gardneri t h a l l u s , t h r o u g h a young monosporangium. Note d i c t y o s o m e s , s m a l l f i b r o u s v e s i c l e s and l a y e r o f f i b r o u s m a t e r i a l b e i n g f o r m e d on t h e o u t s i d e o f t h e plasma membrane . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  168  Transverse s e c t i o n o f Porphyra gardneri t h a l l u s t h r o u g h a monosporangium which h a s d e v e l o p e d beyond t h e s t a g e shown i n F i g u r e 17. S m a l l f i b r o u s v e s i c l e s a r e abundant and l a r g e f i b r o u s v e s i c l e s a r e b e g i n n i n g to form ....................  170  Dictyosome a c t i v i t y vesicles  172  producing  small  fibrous  C l o s e u p o f a m o n o s p o r a n g i a l w a l l showing t h e depositon of fibrous material along i t s inner s u r f a c e (arrow) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  172  Monosporangium i n t h e f i n a l s t a g e s o f d i f f e r e n t i a t i o n p r i o r t o monospore l i b e r a t i o n . B o t h s m a l l and l a r g e f i b r o u s v e s i c l e s a r e abundant  172  Porphyra g a r d n e r i - spermatogenesis, e l e c t r o n microscopy (transverse sections) 20a.  Vegetative c e l l  of the f o l i o s e  thallus  ............  20b.  The f i r s t d i v i s i o n o f t h e spermatangium i s n e a r l y c o m p l e t e . Note t h e new w a l l l a y e r  174  XX {arrow) t h a t h a s been l a i d down i n t h e t r a n s i t i o n from a v e g e t a t i v e c e l l . . . . . . . . . . . . . . . . .  174  Later  174  20c,d. d i v i s i o n s t a g e s o f t h e spermatangium  ........  Porphyra g a r d n e r i - spermatogenesis, e l e c t r o n microscopy {transverse sections) 21a.  F i n a l d i v i s i o n s i n t h e f o r m a t i o n o f a mature s p e r m a t a n g i u m . Note t h e p r o d u c t i o n o f l a r g e Jl l b £ O U S  21b.  21c.  22.  V6 S I C JL*3S  •*•• • • • • • • • •• •  * • ••. •* '*. • • • * • • • • • • • #•-  Spermatium f r o m a mature s p e r m a t a n g i u m . I t c o n t a i n s a nucleus, mitochondria, reduced c h l o r o p l a s t , s e v e r a l s m a l l and one l a r g e f i b r o u s v e s i c l e . Note s m a l l f i b r o u s v e s i c l e emptying i t s c o n t e n t s i n t o t h e l a r g e f i b r o u s v e s i c l e {arrow)  176  Two s p e r m a t i a i n a s p e r m a t a r g i u m n e a r t h e r e l e a s i n g m a r g i n . The l a r g e f i b r o u s v e s i c l e h a s j u s t become e x t r a c y t o p l a s m i c . . . . . . . . . . . . . . . . . . . . . .  176  M a t u r e s p e r m a t i u m j u s t p r i o r t o r e l e a s e . Note t h e n u c l e u s w i t h h i g h l y c o n d e n s e d c h r o m a t i n and no n u c l e a r membrane, r e d u c e d c h l o r o p l a s t w i t h numerous p l a s t o g l o b u l i , m i t o c h o n d r i o n and abundant s m a l l f i b r o u s v e s i c l e s . . . . . . . . . . . . . . . . . . .  178  £oi£kxia. g a r d n e r i - f e r t i l i z a t i o n , Transverse sections 23a.  Spermatium a t t a c h e d the carpogonium  23b. 23c.  1V 6  l i g h t microscopy  t o the prototrichogyne of 180  Two s p e r m a t i a w i t h f e r t i l i z a t i o n c a n a l s g o i n g i n t o t h e carpogonium ..............................  180  Spermatia attached t o both prototrichogynes (note f e r t i l i z a t i o n c a n a l s ) . . . . . . . . . . . . . . . . . . . . . . .  180  Two s c a n n i n g e l e c t r o n m i c r o g r a p h s o f t h e t h a l l u s s u r f a c e o f P o r p h y r a j a r d n e r i showing s e v e r a l spermatia attached to the p r o t o t r i c h o g y n e s ..................................  182  24a,b.  Por£hy.ra g a r d n e r i - f e r t i l i z a t i o n , (transverse sections) 25a.  e l e c t r o n microscopy  Low m a g n i f i c a t i o n v i e w o f a s p e r m a t i u m tc the prototrichogyne  attached  25b-d. T h r e e examples o f s p e r m a t i a a t t a c h e d  to the  184  xxi  prototrichogyne p r i o r t o the formation of the f e r t i l i z a t i o n c a n a l . They c o n t a i n a c h a r a c t e r i s t i c reduced c h l o r o p l a s t , m i t o c h o n d r i a and non-membrane bound n u c l e a r m a t e r i a l .......................................... Porphyra g a r d n e r i 26.  27.  fertilization  T r a n s v e r s e s e c t i o n t h r o u g h a c a r p o g o n i u m and two s p e r m a t i a w h i c h have t r a n s f e r r e d t h e i r n u c l e a r m a t e r i a l down t h e f e r t i l i z a t i o n c a n a l i n t o the carpogonium .............................. Porphyra g a r d n e r i  -  186  fertilization  T r a n s v e r s e s e c t i o n through a carpogonium showing both p r o t o t r i c h o g y n e s , c e n t r a l p y r e n o i d and l a t e r a l n u c l e u s . The p r e s e n c e o f a s m a l l d e p o s i t o f w a l l m a t e r i a l (arrow) i n t h e p r o t o t r i c h o g y n e below t h e s p e r m a t i u m i n d i c a t e s t h a t a f e r t i l i z a t i o n c a n a l h a s been f o r m e d , although i t i s not v i s i b l e i n the plane of s e c t i o n . Suspected Cyanophyta a r e a l s o v i s i b l e on t h e t h a l l u s s u r f a c e ( d o u b l e arrow) . . . . . . . . . . . . . Porphyra gardneri (transverse sections)  184  188  fertilization  28a,b. Two s p e r m a t i a which have t r a n s f e r r e d t h e i r n u c l e a r m a t e r i a l down t h e f e r t i l i z a t i o n c a n a l i n t o t h e c a r p o g o n i u m , but w h i c h s t i l l have t h e r e m a i n s o f a c h l o r o p l a s t l e f t i n them . . . . . . . . . . . . .  190  P r o t o t r i c h o g y n e r e g i o n o f t h e carpogonium with a r e d u c e d c h l o r o p l a s t w h i c h i s t h o u g h t t o have come from t h e s p e r m a t i u m d u r i n g t h e t r a n s f e r o f n u c l e a r m a t e r i a l ..................................  190  28c,d.  Porphyra gardneri 29.  fertilization  T r a n s v e r s e s e c t i o n t h r o u g h a carpogonium which has t h e r e m a i n s o f a s p e r m a t i u m a t t a c h e d t o t h e p r o t o t r i c h o g y n e , and what may be t h e h i g h l y c o n d e n s e d s p e r m a t i a l n u c l e a r m a t e r i a l (arrow) .....  192  Porphyra gardneri - carpcsporogenesis (transverse sections) 30a.  F e r t i l i z e d c a r p o g o n i u m s h o w i n g t h e two d a u g h t e r n u c l e i r e s u l t i n g from t h e f i r s t d i v i s i o n o f t h e f u s i o n n u c l e u s . The f e r t i l i z a t i o n c a n a l i s s t i l l p r e s e n t a l t h o u g h t h e spermatium has d i s i n t e g r a t e d ................ . . . . . . . . . . . . . . . . . . . . . . . .  194  xxii  30b.  The f i r s t d i v i s i o n o f t h e c a r p o s p o r a n g i u m i s p e r i c l i n a l and produces 2 c a r p o s p o r e s (note t h e f e r t i l i z a t i o n c a n a l s t i l l p r e s e n t i n t h e w a l l ) ..... 194 P2rj3hy_ra  31a.  Spermatium,  31b.  Vegetative  g a r d n e r i - chromosome c o u n t s  n=4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . cell  of t h e f o l i o s e  thallus,  n=4 .......  196 196  31c,d. C a r p o s p o r e , 2n=8  196  31e,f. Vegetative  cell  of the conchocelis  Porphyra g a r d n e r i - Feulgen (transverse sections) 32a. 32b.  32c.  33b.  196  staining  Mature s p e r m a t a n g i u m , most s p e r m a t i a s t i l l have n u c l e i i n a d i f f u s e i n t e r p h a s e s t a t e (arrow) ......  198  M a t u r e s p e r m a t a n g i u m r i g h t on t h e r e l e a s i n g margin. The n u c l e a r m a t e r i a l i s h i g h l y c o n d e n s e d (arrow) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  198  Seleased spermatia with highly n u c l e a r m a t e r i a l (dark d o t s )  198  condensed  Porphyra g a r d n e r i - Feulgen (transverse sections) 33a.  phase, 2n=8 ....  staining  C a r p o g o n i u m and a t t a c h e d s p e r m a t i u m w h i c h c o n t a i n s c o n d e n s e d n u c l e a r m a t e r i a l . No f e r t i l i z a t i o n c a n a l i s p r e s e n t ....................  200  Carpogcnium and c a r p o g o n i a l nucleus l a t e r a l l y l o c a t e d . The a t t a c h e d s p e r m a t i u m c o n t a i n s n u c l e a r m a t e r i a l i n a d i f f u s e s t a t e , and a f e r t i l i z a t i o n c a n a l i s p r e s e n t ....................  200  Two e x a m p l e s o f s p e r m a t i a c o n t a i n i n g c o n d e n s e d nuclear m a t e r i a l i n c l o s e a s s o c i a t i o n with the f e r t i l i z a t i o n c a n a l ...............................  200  Carpogonium w i t h a t t a c h e d empty s p e r m a t i u m and a d e n s e l y s t a i n i n g mass o f s p e r m a t i a l n u c l e a r m a t e r i a l i n the carpogonium c l o s e to the f e r t i l i z a t i o n c a n a l ...............................  200  S e v e r a l examples o f c a r p o g o n i a w i t h empty s p e r m a t i a on t h e i r o u t e r w a l l s and from 1-5 d e n s e l y s t a i n i n g masses o f s p e r m a t i a l n u c l e a r material inside  200  33c,d.  33e.  33f-i.  xxiii  33j.  33k. 331.  Carpogonium with l a r g e , s u s p e c t e d f u s i o n n u c l e u s (note empty s p e r m a t i u m and f e r t i l i z a t i o n canal) ..............................  200  The f i r s t d i v i s i o n o f t h e f u s i o n n u c l e u s p r i o r t o c y t o k i n e s i s (see a l s o F i g u r e 30a) . . . . . . . . . . . . . .  200  Two c a r p o s p o r e s p r o d u c e d by t h e f i r s t d i v i s i o n o f t h e c a r p o s p o r a n g i u m ( s e e a l s o F i g u r e 30b) ......  200  Subseguent s t a g e s o f d i v i s i o n t o produce 4 ( F i g u r e 33m) o r 8 ( F i g u r e 33o) c a r p o s p o r e s ........  200  D i a g r a m s u m m a r i z i n g t h e known m o r p h o l o g i c a l and c y t o l o g i c a l l i f e h i s t o r y c f Porphyra gardneri. I t i s suspected that meiosis occurs i n t h e conchosporangium at t h e t i m e o f conchospore f o r m a t i o n .........................................  202  Porphyra n e r e o c y s t i s spermatium i n a s p e r m a t a n g i u m n e a r t h e r e l e a s i n g m a r g i n . The s p e r m a t i u m c o n t a i n s a b u n d a n t s m a l l and l a r g e f i b r o u s v e s i c l e s , a reduced chloroplast, m i t o c h o n d r i a , and c e n t r a l n u c l e u s . . . . . . . . . . . . . . . . .  204  33tt-o.  34.  35.  Porphyra  nereocystis  36a,b.  36c.  Transverse s e c t i o n of the f o l i o s e t h a l l u s t h r o u g h c a r p o g o n i a and s u s p e c t e d s p e r m a t i a forming f e r t i l i z a t i o n c a n a l s ......................  206  Electron micrograph o f a transverse s e c t i o n t h r o u g h a s u s p e c t e d s p e r m a t i u m and f e r t i l i z a t i o n c a n a l ...............................  206  Porphyra 36d. 36e.  thuretii  T r a n s v e r s e s e c t i o n of t h e f o l i o s e t h a l l u s t h r o u g h a c a r p o g o n i u m and s u s p e c t e d s p e r m a t i u m  ...., 206  Transverse section of the f o l i o s e t h a l l u s t h r o u g h a c a r p o s p o r a n g i u m t h a t h a s undergone t h e f i r s t d i v i s i o n . A s u s p e c t e d s p e r m a t i u m and f e r t i l i z a t i o n c a n a l a r e p r e s e n t ...................  206  37a.  Porphyra n e r e o c y s t i s  spermatia,  208  37b.  Porphyra nereocystis  vegetative c e l l s  foliose  ..............................  208  P o r p h y r a t h u r e t i i s p e r m a t i u m , n=2 . . . . . . . . . . . . . . . . .  208  37d,e P o r p h y r a n e r e o c y s t i s c a r p o s p o r e , 2n=6. ( o n l y 5 chromosomes v i s i b l e ) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  208  37c.  thallus,  n=3  n=3 . . . . . . . . . . . . . . . of the  xx i v  37f,g P o r p h y r a 2n=6  .7  nereocystis  conchosporangial branch, . . ......  208  XXV L I S T OF  I. II.  APPENDICES  P o r p h y r a c o l l e c t i n g s t a t i o n s and d a t e s o f c o l l e c t i o n s ........................................  234  R e p r e s e n t a t i v e specimens of Porphyra g a r d n e r i , P. n e r e o c y s t i s and P. t h u r e t i i . . . . . . . . . . . . . . . . . . . . .  236  xxvi  ACKNOWLEDGEMENTS I  am  indebted  facilities, A-4471)  advice  t o D r . fl. F. S c a g e l  and g e n e r o u s f i n a n c i a l  throughout  this  study.  Canada P o s t g r a d u a t e s c h o l a r s h i p s this  work.  providing  Dr. K, C o l e  my  to  specimens  express  information Dr.  fungal  gratefully  forgotten.  The  Thanks  introducing  me  to  I . A. A b b o t t  R.  specimens.  excellent  are  given  for  also  the  stay  holotype  of  specimens from  Porphyrella to  of  Porphyra.  with  E.  Foreman and D r . in  some  to  me  the  of  the P.  manuscript. assistance  comradeship  at  which  with  Hopkins  gardneri:  laszlo  t o David  excellent  herbarium for  to  i s  Walker f o r technique.  Station,  and  specimens i n h e r lending  and for  Veto  f a c i l i t i e s and  Marine  californica  Amchitka I s l a n d ;  by  on e l e c t r o n  methacrylate  Dr. P. C. S i l v a  Porphyrella  advice  extended  JB-4 me  me  Thank you t o Dr. G. J . H o l l e n b e r g  gardneri;  in  D r . G. C. Hughes  The  to  me t o examine G. M. S m i t h  Porphyra  used  would  technical  permitted  of  for  t o t h e s e a and t o t h e m o u n t a i n s w i l l n o t  during  holotype  acknowledged  Tanner f o r h i s u n f a i l i n g  assistance  care.  out  I  improvements  provided my  providing  and t o D r .  and p h o t o g r a p h y  appreciated.  to  Grant  carry  equipment  infections  for  to collect  on o u r t r i p s  me t o  forthe cytological studies.  f o r suggesting  SCOBA d i v e s  (NEC  Research C o u n c i l of  have e n a b l e d  i s  t h a n k s go t o C h r i s  microscopy  Dr.  National  appreciation  literature,  developed be  on  Harrison  Special on  my  T. F. Mumford J r . ,  pertinant G.  support  me w i t h a c c e s s t o h e r l a b o r a t o r y  preparing like  f o r use o f t h e c u l t u r e  the  me  the  specimens  of  loan  of  Dr. M. J . Wynne  and t o R. S e t z e r  for  the  the for loan  xxvii  of  specimens o f Porphyra My  The  parents i n t e r e s t  encouragement  made t h e  and  gardneri. i n my  patient  t a s k of w r i t i n g  s t u d i e s has understanding  this thesis  been an  inspiration.  of Denise  tolerable.  Bonin  has  1  GENERAL INTRODUCTION  Despite Porphyra the is  the  g r e a t amount o f i n v e s t i g a t i o n  has r e c e i v e d ,  complete  life  Higita's  most  thorough  the  present a d e t a i l e d which  is  the  reguiring One this  the  b o t h m o r p h o l o g i c a l and  (1967b) work on study  to  date.  documentation  major  Porphyra  cytological,  y e z o g n s i s Ueda  U n f o r t u n a t e l y he  of the f e r t i l i z a t i o n  feature  of  the  which  Porphyra  process,  life  history  to  rectify  verification. o f my  primary o b j e c t i v e s  situation  by  attempting  o t h e r main o b j e c t i v e  similar  epiphytic,  as  information  much  b i o l o g y as  i n t h i s s t u d y was to  answer  the  much d i s p u t e d  study  third  was  approach for  indicating light  and  Porphyra life  to t h e s e  3  superficially  s p e c i e s by histories  obtaining and  a l l  any  basic  of spore formation.  on s p o r e  spore in  occurrence  electron  microscope  study life  ploidy of  to  first  of  second  of  obtain  the was  selected  germination.  The  chromosome  history  stages to see i f  level,  thus  fertilization study  The  the e f f e c t  f o r m a t i o n and  t y p e s and the  The  the s e a s o n a l o c c u r r e n c e  a cytological  change  the  p r o b l e m s were t a k e n .  o f s p o r e f o r m a t i o n and factors  t h e r e was  characterize  their  study t o determine  environmental  counts  about  p h a s e and t h e t i m e  a culture  to  monostromatic  approaches  a field  foliose  was  genus.  possible.  Three was  is  d i d not  g u e s t i o n of t h e o c c u r r e n c e o f s e x u a l r e p r o d u c t i o n i n the The  genus  t h e r e i s not a s i n g l e s p e c i e s f o r  history,  known.  that  of  and  indirectly meiosis.  sporogenesis  by  A the  2  foliose  thallus  give d i r e c t  evidence  Initially Anderson.  I  It  r e p o r t e d l y very be  a  habit  plant also  special its  was  undertaken  o f the occurrence  began  is  working  of  with  that  i t might  fertilization. Porphyra  nereocystis  one o f t h e l a r g e s t Porphyra s p e c i e s ,  palatable  with  because I f e l t  (Hus, 1902).  potential  I t t h e r e f o r e seemed  commercial v a l u e .  i n t r i g u e d me, and  I  was  and i s  curious  I t s epiphytic  to  see  i f any  l i f e h i s t o r y m o d i f i c a t i o n s had been made t o a d a p t  •host*  plant,  Nereocystis  luetkeana  to  i t to  (Mertens) P o s t e l s e t  Buprecht. During of  the f i r s t  year  Porphyra g a r d n e r i  superficially considered  (Smith  resembles  by  o f my f i e l d  Gardner  et  studies I  Hollenberg)  Porphyra n e r e o c y s t i s (in: Collins  et  (1941) t o be a dwarf o r young f o r m o f t h i s this,  and  because  published  on  Hollenberg  (1943),  was  a  fortunate  convenient this  this  life  alga  history  since  During  This  which  and  species.  information  i t was d e s c r i b e d  because  Porphyra  second  year  was  had  i t proved  I n view o f had  been  by S m i t h and study.  This  t o be t h e most  As a r e s u l t , t h e major p a r t  of  gardneri. of  my  field  observations  S e t c h e l l e t Dawson on t h e s t i p e  the  been  a l . 1919) and K y l i n  decision  Porphyria t h u r e t i i  M§£§°cy.stis.  Hawkes*,  t o i n c l u d e i t i n my  with  the  aware  I decided  p l a n t t o work w i t h .  thesis deals  collected  no  became  first  time that  this  I of  s p e c i e s had  lhen I began my investigation, this plant was known a s P o r p h y r e l l a g a r d n e r i Smith e t Hollenberg. As a result o f my observations o f t h e mode o f c a r p o g o n i u m f o r m a t i o n and d i v i s i o n (to be d i s c u s s e d i n P a r t I I ) I t r a n s f e r r e d t h i s s p e c i e s t o t h e genus P o r p h y r a (Hawkes, 1977b). l  3  been f o u n d little  was  north  known  a l g a I decided I and  have p r e s e n t e d herbarium  Cytological studies the  transfer  (Conway e t  about  to include  carposporogenesis;  of  o f Oregon  the  the  ecology  i t i n ay  and  Part  II  I I i s one  which I have p r e s e n t e d because  it  so this  aspect  Field  genus  and  Part  and IV-  of the c y t o l o g i c a l  near  essential  of P o r p h y r e l l a aardjneri t o t h e  I -  Spermatogenesis  studies;  separately, is  h i s t o r y of  parts: Part -  Part I I I - Culture Part  life  Because  study.  thesis i n four  studies;  studies.  thesis,  a l . 1975).  the  beginning  in justifying Porphyra.  the  4  TERMINOLOGY  The  morphological l i f e  consists  of  an  alternation  f i l a m e n t o u s phase. stages  as  phase,  the  I will  foliose  of  the  between  be r e f e r r i n g thallus  i s a plethora  reproductive  work on  the  of  structures  to these  life  and  a  history  and t h e c o n c h o c e l i s  Bangiophycidae  Drew  (1956)  a scheme s p e c i f i c a l l y  As a r e s u l t  o f my  sexual  reproduction  Porphyra  gardneri.  terminology  was  a  which  occurs  a detailed  f o r the  as  well  as  other  reproductive (1975,  Foliose  summary  The  In her  classic  presented  Porphyra.  the  a  spore  Richardson Porphyra. 1978)  life  and  that  history  of  standardization  of  following  reproductive  i s a summary c f t h e  structures  and  spore  i n t h e 3 Porph,yra s p e c i e s I s t u d i e d .  of the o r i g i n a l  terminology  s t r u c t u r e s o f the  p.  applied  f o r B a n g i a and  in  reassessment  which I e n c o u n t e r e d  For  included  been  o b s e r v a t i o n (Hawkes, 1977a,  necessary.  I have used  has  of Porphyra.  (1972) p r o p o s e d  al.  species  thallus  t e r m i n o l o g y which  scheme  types  Porphyra  a foliose  o r phase,  classification  terms  most  respectively.  There to  history  that foliose  sources of has  these  terms  been a p p l i e d  t o the  phase  see  Conway  et  a  series  of  189) .  Phase  Spermatangium  -  produces  periclinal  and  the  spermatia  anticlinal  by  divisions.  Synonymous  5  with  J2> - s p o r e  packet  (Hus,  Kurogi,  Spermatium  -  the  -  gamete.  by  may  or  Synonymous  Hawkes,  -  may  mother  varies  carpogonium,  and  with  antherzoids  transformation not  produce  cells  of a  vegetative  prototrichogynes.  cells  (Drew,  (Conway  1956)  et  and  a l . 1975;  protuberance of the  with the s p e c i e s  carpogonium.  and n o t a l l s p e c i e s  Honcstromatic species whereas d i s t r o m a t i c  p r o d u c e 2 per  species  produce  one.  -  following  becomes  the  carpospores anticlinal  fertilization  the  carposporangium. by  a  divisions.  series The  of  packet,  sporocarps  the  periclinal  and  possibility  (Hus,  1 9 7 2 ) , and p a c k e t s o f  a l . 1975;  Hawkes,  1977b).  carpogonium  Produces  can o c c u r w i t h o u t f e r t i l i z a t i o n .  (Kurogi, et  1902;  1977b).  p r o d u c e them.  with  (Hus,  Synonymous  a l . 1975),  mother  receptive  Length  this  the  with  -spore  Carposporangium  1975),  1902).  cell,  only  antheridium  Haploid.  (Conway e t  formed  lEototrichogyne  (Conway e t a l .  1972).  ^-spore  Cargogonium  cell  1902), ana  male  (Hus,  mother  1902),  exists  that  Synonymous cystocarps  O^-spores  (Conway  6  Carpospore  -  released into  the  haploid  from t h e carposporangium conchocelis  phase.  carpospores  may  Syncnymous w i t h Hawkes,  Monos£ore  -  several  spore  Following  produced  cell  release  foliose  phase.  (eg. Kurogi,  In  referred  to  of  1953b) as  by  phase  and  1975;  phase o f  divisions.  the c o n c h o c e l i s  phase  the  the  which  into spore  Japanese  conchospores  monospores.  been a p p l i e d  more c o n c h o c e l i s  any  some  monospore has a l s o  is  al.  Synonymous w i t h n e u t r a l  literature  QgSghoseorangial  cases.  a monospore g e r m i n a t e s back  1961).  Conchocelis  gt  by the f o l i o s e  without  (Kurogi,  were  (Conway  some  P o r p h y r a s p e c i e s by t h e t r a n s f o r m a t i o n o f  vegetative  the  in  although  1977b).  asexual  a  Diploid,  form  ^-spores  and g e r m i n a t e s  The  t o spores germinate  (Conway a n d C o l e ,  term  produced to  give  1977).  Phase  branch - t y p i c a l l y  an u n i s e r i a t e b r a n c h  wider than t h e v e g e t a t i v e c o n c h o c e l i s consists  of  conchospor angia. w a l l compared  linear  Characterized  to the vegetative  conchosporangium chloroplast.  a  contains  The a v a i l a b l e  a  filaments  series by  a thick  filaments. single  which  of cell Each  stellate  evidence suggests that  7  the  conchosporangium  (Giraud  and  in  1967b;  1974).  some  some  of  {Kurogi,  (Migita  1974). with  synonymous 1953b).  Abe,  1966;  not c l e a r  Referred  Japanese  meiosis  Kito,  Migita,  whether  per conchosporangium  species. the  of  o f the conchosporangium  and  It i s s t i l l  one c o n c h o s p o r e by  is  by 2-4 d i v i s i o n s  some s p e c i e s  site  1968;  branch  monosporangial branch  - formed  the  Magne,  Conchosporangial  Conchospores  i s  is  only  produced  t o a s monospores i n  literature  (eg. K u r o g i ,  1953b) . A  brief  justification  of  r e p r o d u c t i v e s t r u c t u r e s formed order. of  Functionally  my s e l e c t i o n  by  the  foliose  consistent  with  standardize spermatium, been  I n view  previous  of this,  work  i s in  spermatangium,  counterparts  in  and i n an a t t e m p t t o be  (Papenfuss,  t h e t e r m i n o l o g y i n the two  1955),  subclasses,  carpogonium  and  and  the  to  terms  c a r p o s p o r e s have  used. Differences i n the position  spermatia  and  carpospores  characterizing (Fritsch, spermatia only  phase  t h e s p e r m a t i a , c a r p o g o n i a and c a r p o s p o r e s  P o r p h y r a g a r d n e r i a r e homologous w i t h t h e i r  the F l o r i d e o p h y c i d a e .  of terms f o r the  1945,  the  one i s p r o d u c e d .  been  Bangiophycidae  p. 437;  are produced  have  and mode o f f o r m a t i o n  Drew,  pointed and  1951).  of  the  t o as c r i t e r i a  Florideophycidae  In t h e former, s e v e r a l  p e r s p e r n a t a n g i u m , whereas i n t h e l a t t e r After  the e n t i r e zygote d i v i d e s  fertilization  t o produce  i n Porphyra  several  gardneri  carpospores  per  8  carposporangium, directly  should  really  be  noted  For  spermatangium  to  is  the  carpospore  the d e t a i l s  example,  and  It  are not o f s u f f i c i e n t  zygote  gonimoblast per  production  apparently o n l y one  is  whose  carposporangium.  of spermatium  i n Smithora  produced.  the  felt  not  a l l  spermatium  that  magnitude t o warrant  these creating  terms f o r t h e r e p r o d u c t i v e s t r u c t u r e s i n P o r p h y r a . The  species  slight cf  protuberances  Porphyra  pseudo-trichogynes and Chang,  1955),  trichogynes  receptive  (Table  VI)  1  f o r the  the s p e c i f i c i t y homologous  however, b e c a u s e prctctrichogyne  have  1927),  trichogyne-like* 1939).  protuberances  site  of t h e carpogonium  (Dangeard,  (Kunieda,  specialized  are  rise  a single that  Florideophycidae  only apply t o t h e Bangiaceae,  differences new  the  gives  produce  Bangiophycidae. per  in  or i n d i r e c t l y  f i l a m e n t s then It  but  of  with  the  they  are  referred  to  prototrichogynes  (Conway and Porphyra  the  carpogonium seem t o be  attachment.  gardneri act  as  (Tseng  C o l e , 1973)  In  s p e r m a t i a and  o f spermatium  been  f o u n d i n most  and these  as  the  responsible for  Functionally  they  t r i c h o g y n e s of the F l o r i d e o p h y c i d a e ; not  as  well  developed  would seem most a p p r o p r i a t e .  the  term  9  PART I - FIELD AND HERBARIUM  STUDIES  Introduction Before  attempting  investigation familiarize £•  myself  thuretii  to  I felt  each fix  chromosome  at  the  numbers  these  3  or  utmost  cytological  importance  of the f i e l d  study  s e a s o n a l o c c u r r e n c e , t y p e and t i m e geographical  I hoped t h a t t h i s  thalli  work  right  and  time  distribution  for cytological reproduction.  are  was  of spore  i n f o r m a t i o n would e n a b l e  sexual  species  to  aardjae_ri, £.. n e r e o c y s t i s . and  The p u r p o s e  * h o s t ' s p e c i e s , and  species.  because  Poo^lia  with  the  culture  t h a t i t was o f t h e  i n the f i e l d .  determine  formation,  any  for me t o  study  of  Furthermore,  e p i p h y t e s , I thought  t h a t some  knowledge o f t h e s e a s o n a l d e v e l o p m e n t o f t h e • host{s)« would be essential The  t o a complete  following  understanding  i s a brief  of their  characterization  life  histories.  o f t h e 3 s p e c i e s and  a summary o f p r e v i o u s i n v e s t i g a t i o n s o f them. Porphyra which  grows  Laminariales Smith  gardneri i s a epiphytically {Figure 1a).  Peninsula,  California By  tentatively  called  spermatia Porpjjxra,  but  the  from  were  late  were s a i d  several  In the o r i g i n a l  summer t h a l l i  monospores.  on  monostromatic  (1943, as P o r p h yr e l i a  and H o l l e n b e r g  that i n early  foliose,  Point  either  summer  carpospores to  occur  carpospores  ga i d p e r i on  and being  packets  of  description  as  the  ) reported  or  Monterey producing  spores  they  produced. i s  alga  of i t .  the  vegetative  were  in  members  Joe  spermatia  red  The  typical  were r e p o r t e d t o o c c u r  of  singly.  10  Smith  and H o l l e n b e r g  formed  by a c e l l  guite  (1943, p. -.215) s t a t e d , "The c a r p o g o n i a a r e  division  similar  to  i n which t h e r e  the  curving  cell  moncspores o f E r y t h r o t r i c h i a  and  liberated  single  cell  singly. '  These  1  division  were  the  the  basis  i s  cells  spores  the  formed  are  formed by an u n e q u a l  f o r establishing  type  species  ( h o l o t y p e , G. M. S m i t h 39-12, DS  The  fate  spores  wall,  producing  thus  (Bangiophycidae),  these  curving  walls  forphyrella  of  a  the  genus  with P o r p h y r e l l a g a r d n e r i as the  was  not  306401  in  UC) .  d e t e r m i n e d by S m i t h and  Hollenberg, Subsequent  workers  have  Porphyra j a r d n e r i  (as  California  1944;  Oregon and  (Smith,  (Doty,  British  Alaska  1947;  Columbia  (Wynne,  information  the  occurrence  Porphyrella gardneri) Abbott  and  1972;  of  elsewhere  Hollenberg,  1976),  Markham and C e l e s t i n o , 1976),  (Scagel,  concerning  reported  the l i f e  1977),  history  in  Washington  1957, 1973; Widdowscn, 1974)  Lindstrcm,  in  and  However, no f u r t h e r  of this alga  has  been  published. Porphyra  nereocystis  Bangiophycidae  which  Nereocystis luetkeana of  the l a r g e s t  i s  grows  a  monostroaatic  epiphytically  (Mertens)  3,7 i i n l e n g t h a n d 0.7 m i n w i d t h Vancouver  Island).  name o n l y ;  description published  1892)  based  Islands, that  on  California.  Pyropia  from  No h o l o t y p e  californica  the  ofthe  stipe  from  Blankinship  Santa  Cruz  exists.  J . Agardh  or Hus  i s  a  up t o  Esguimalt,  r e p o r t e d by Anderson  in  of  I t i s one  (Figure 1b), reaching (UC 96517,  was f i r s t  material  on  P o s t e l s e t Buprecht.  species of Porphyra  It  member  and  (1891, Keeler,  the Farallon (1900)  noted  synonym  of  11  22IEhl£.* n e r e o c y s t i s .  has  Porphyra n e r e o c y s t i s i s the  only  been  w o r k e r s t o any  i n v e s t i g a t e d by  been  reported  1944;  Abbott  Washington  in California and  1925;  (Collins,  Conway e t a l . , 1 975)  1903;  J o h a n s e n , 1971; (1902) was  nereocystis  commercial  al.  (1969a),  and  first  i t s use of  investigation  conchocelis Mumford  (1977).  Alaska  by  the  which  extent.  Kylin,  Oregon  Scagel,  to  by  It  1941*;  has  Smith,  {Doty,  1947),  1972;  Mumford,  1973a),  1957,  1973;  (Setchell  wlddcwson,  and  Gardner,  1977). comment on the  the  palatability  Chinese i n C a l i f o r n i a .  plant  has  Woessner  t h a l l u s and Despite this  recently  (1974)  of  reports of  seldom  until  Nereocystis  the  is still  and  found  Cole  and  are  not  of The  received Woessner e t  past well  cell  Krishnamurthy  (1?75) and  Conway  reported  of  both  on  the  and the  foliose  phase. amount o f s t u d y , w e l l known.  i t s seasonal  Keeler,  by  (1975) b r i e f l y  vegetative  a considerable  Blankinship  been c u l t u r e d  ( 1 9 7 3 a ) , Conway e t a l .  the c o n c h o c e l i s  alga  conflicting  phase h a s  Conway and  ultrastructure  of  1913;  studied  (1977). The  Cole  and  potential  extensive  1976),  Lindstrom,  the  1893;  Krishnamurtby,  1974;  P.  (Howe,  Hollenberg,  {Kylin,  B r i t i s h Columbia  Hus  other  species I  1892)  life  history  For example, t h e r e  occurrence.  Anderson  only i n d i c a t e d that i t  midsummer, grown."  the  when  Smith  * K y l i n mistakenly r e f e r r e d specimens Porphyra n e r e o c y s t i s  of  the  are (in  "...is  long  stems  of  and  Abbott  and  Porphyra g a r d n e r i  to  (1944)  12  Hollenberg and  is  al.  (1976) r e p o r t e d  usually  (1977) and  species. cf  found  In contrast, as  the f o l i o s e  phase i s an  from November t o J u n e .  Arasaki  occurrence  that  (1974)  agree  that  Both  Hoessner e t  i t i s  Conway e £ al,. (1975) r e p o r t e d  summer  through  winter,  annual  a  winter  t h e season  and Conway and C o l e  (1977) i n d i c a t e d i t i s summer. There are also specificity  exhibited  Blankinship  and  frequently  found  United  to i t .  Abbott  and  by  Keeler, on Hus  (1902)  that  the  Peninsula not was the  (on  little  (1944) species  included  which was i t  California,  P a c i f i c Grove.  according herbarium,  most  exclusively on  rocks.  i t i s occasionaly  (1974)  felt  i t  was  and  the to  was  particularly  occurrence.  Paul  to the lectotype,  published  actually in  epiphytic  flaring  in  published Algae  Of  but s t a t e d t h a t  Dawson  label  the  (1944) d e s i g n a t e d  C.  i t s Silva  10 D e c ,  was  intended  July  1944.  first  by S m i t h  The  Monterey  which a p p e a r e d e a r l y i n 1944.  i n d i c a t i n g that to  known,  p u b l i c a t i o n o f P. t h u r e t i i  c i t e a type c o l l e c t i o n  type,  not  monpstromatic,  i s  a note attached  the o r i g i n a l  who  another  plants or i t s seasonal  be i n Dawson  (1944)  that  (in  i t was  i t growing  but Soessner  is  which v e r y  indicated  However,  reported  host  Anderson  that  although  of  to Nereocystis.  range o f host  to  indicated  (1976) i n d i c a t e d  Laminariales,  P o r p h y r a about  1971)  o f the degree  P. n e r e o c y s t i s .  1892)  Porphyra t h u r e t i i  has  reports  Nereocystis.  Hollenberg  f o u n d on o t h e r exclusive  conflicting  type  Smith d i d locality  S e t c h e l l #5161 as  i t was from P a c i f i c  Grove;  on s p e c i m e n s o f t h a t  number i n t h e UC  Setchell's  field  book,  his  however,  #5161  was  13  collected  at  collection  (DC 791973)  P. C. S i l v a growing  Carmel  in  Bay.  A  was  1949  i n addition  was  from  known  reported  this  this  as  (Figure  on G r a c i l a r i a  (1944) n o t e d t h a t also  from  designated  September  epiphytically  plant  the  1c).  lectotype It  was  by  found  sjoestedtii Kylin.  Dawson  t o the type l o c a l i t y , t h e s p e c i e s  S a n t a C r u z and Monterey  species  29 Hay 1900  under  the  Bay.  Hus  (1902)  name  Porphyra  leucpsticta  subsequently  reported  from  Thuret. Porphyra t h u r e t i i (Doty,  1947).  Columbia  It  was  has  f o rthe f i r s t  recently  time  I made a t my s t u d y s i t e  Island.  Dawson  l o c a t i o n s i n Mexico, (1945)  and  collected  in  Both  Smith  P. t h u r e t i i Da«scn  (1944)  grows  (1952) or  et  (1975)  j£ieoc£stis The  rarely  that  Bica  and  that  Dawson  the  epiphytic.  was  Sound,  Vancouver  plants  reported  sjoestedtii  on  Taylor to  as  P. t h u r e t i i .  from  or  on  Mexico  The p l a n t r e p o r t e d  growing  several  referred  (1944)  plants  British  from  the  and  , were a c t u a l l y  in  1975), b a s e d on a  P. t h u r e t i i  indicated  Gracilaria  stated  saxicolous a l .  on  i n Barkley  reported  Costa  P o r p h y r a naiadum A n d e r s e n  reported  ( Conway e t a l .  1  collection  (1952)  been  Oregon  the  that rocks. were  by Conway  stipe  of  luetkeana.  season  (March t o May)  o f o c c u r r e n c e has been r e p o r t e d  (Hus,  1902,  as  P. l e u c o s t i c t a ;  as t h e s p r i n g Smith,  1944;  *Scagel (1973) r e c o r d e d P. t h u r e t i i f r o m B a r k l e y Sound, b u t Conway e t a l . (1975) indicated that these specimens are probably more correctly identified as Porphyra abbottae K r i s h n a m u r t h y a n d P. f u c i c o l a K r i s h n a m u r t h y .  14  Dawson, 1944),  and summer  ( Conway a n d C o l e  Materials Most Marine  of  Station  Columbia  the f i e l d  A study  (48° 5 0 . 1 » N ,  because  i t i s semi-exposed  •Zodiac* At °  channel  landed  And Methods  work was c a r r i e d  Sound  be  site  out near  was  most  inflatable  conditions  boat  t h e study s i t e  specimens c o l l e c t e d  was u s e d  this  British  i n Barkley  selected  ( F i g u r e 2b)  Ocean and has  Island,  a  where a b o a t c a n  throughout  fora l l field  f . g a r d n e r i reached  from  Island,  t o t h e open P a c i f i c  between i t and D i a n a  under  the Bamfield  o f f Diana I s l a n d  125° 11.1*w)  Laminaria s e t c h e l l i i S i l v a ,  n  1977).  on t h e west c o a s t o f V a n c o u v e r  (Figure 2a).  sheltered  ,  the  year.  A  work.  optimal  development  Most o b s e r v a t i o n s were made on  *host*  from  May 1974 t h r o u g h  May  1977. Other collected Cape  sites  were: Ross I s l e t s ,  Beale.  collecting  See  at  site  the  for a  location,  Barkley  Islet.  from J u n e  and  Execution complete  and d a t e s  Sound  the kelp beds o f f : A g u i l a r At t h e s t u d y  1974 t h r o u g h  RQLSklE.!* t h u r e t i i 1975  I  Islet,  Rock, list  and  of  my  of c o l l e c t i o n .  y e a r t o y e a r s o i t was n e c e s s a r y t o c o l l e c t i t  in  Beach, and L e a c h observed  their  Leach  g a r d n e r i was  n e r e o c y s t i s was v a r i a b l e i n i t s abundance a t t h e  from  other s i t e s  included  Appendix  stations,  Porphyra study  i n B a r k l e y Sound where P o r p h y r a  April  area  as  Point, Cable site  well.  These  Beach,  Second  P. n e r e o c y s t i s  was  1977.,  was n o t o b s e r v e d  a t t h e study  was n e v e r a b u n d a n t e n o u g h t o make a d e t a i l e d  site  until  monthly  15  study at  of  i t .  Recently  Leach I s l e t , Porphyra  or l e s s ) ,  !§I§2CV.stis proved  gardneri  at  all  hosts  the  for  could  making  P.  SCUBA was in  reached only  by  the  opportunity In  B r i t i s h Columbia: Islands,  Peninsula;  California:  Ano  Monterey  Langara  the  collected Charlotte Charlotte  Barkley included:  Islands;  Pender  Island.  area  British  K y u g u o t Sound; B o t a n i c a l  Islands;  Point  Mission  Sound  Tree  Queen  Renfrew.  and  where  electron subtidal was  nereocystis conchocelis  the  study  site,  Joe  Islets;  Brooks  Cove;  and  Harris  Beach.  Pescadero  Point,  Carmel.  Porphyra  Localities  nereocystis  Langara  Skincuttle Hope I s l a n d ;  Beach; W h i f f e n Pebble  Springs  were:  Islands;  Tree I s l e t s ;  and  Point,  these  Charlotte  Oregon:  Columbia:  Islands,  California:  and and  SCUEA  for  at  Kyuguot Sound; Hot  Bolkus  Islands;  chamber  a  s p e c i m e n s were c o l l e c t e d f r o m  Island,  Nuevo I s l a n d ;  Peninsula;  artificial  made  Charlotte  Port  used  several SCUBA.  be up  observations  using  m  could  pulling  case c f Porphyra gardneri  Grassy I s l a n d , Beach,  by  (0.5  measured.  arose,  the  Queen  Botanical  outside  were  s e a w a t e r t e m p e r a t u r e was  localities.  Gordon  culture  When c o l l e c t i o n s  Hhen other  situ  Sound.  tides  freguently  a l s o grows on  an  surface  abundance  thuretii  situ  plants  in  low  t i d e s simply  anchor experiments u s i n g  filaments.  in  f o r chromosome c o u n t s and  gardneri be  and  a l s o needed to and  found  c o l l e c t e d on  this,  material  Porphyra  be  highest  Despite  fresh  which  i t was  nereocystis  but  invaluable  microscopy.  could  P.  plant.  collecting  1977)  a much more e x p o s e d l o c a t i o n i n B a r k l e y  whereas  obtained  (April  was  Island,  Queen  Inlet,  Queen  Grassy  S p i t , Sooke; and  Beach, M o n t e r e y  Island, South  Peninsula.  16  Other s i t e s collected  i n British  Columbia  included: Bennell  where  Porphyra  thuretii  Sound, Queen C h a r l o t t e I s l a n d s and  V o l c a n i c C o v e , Kyuguot C h a n n e l , V a n c o u v e r I s l a n d . of  observing  population The  field  these  other  sites  The  purpose  was t o s e e i f t h e r e  s e r e any  d i f f e r e n c e s i n t h e t i m e and t y p e study  of  Porphyra gardneri  Monterey  {Point  Joe,  confirm  t h a t t h e p l a n t s a t the study  at t h e s t u d y Islands, several  where  £•  of  the  the stipe.  taken  in  A series of a r t i f i c i a l was  the  the  environment  Three  types  manila  rope  brand, brand, *stipe*  host  floatation.  host  Charlotte  (3-strand,  15 mm  outside  lashed The  to  a  and  of  A l l the  taken.  requires.  s t i p e s were made, u s i n g vinyl  tubing  (Fisher  rubber  (Fisher  A t one end t h e  plastic  anchored  bottle  to rock  for  climbing  pitons  (Cassin - I t a l y ,  ring  47 mm  diameter)  which  were d r i v e n i n t o c r e v i c e s i n t h e r o c k y  bottom  a t a depth  of  7-10 m.  Polypropylene  rope  long with  position i n  epiphyte  surgical  sealed  ' p l a n t s * were t h e n 250 mm  a  pneumatocyst,  forthe 'stipe*.  4 1  get  distribution  the unique  that the  diameter),  diameter)  to  e x p e r i m e n t s was r u n t o s e e i f  Nereocystis  diameter)  order  meter were c o l l e c t e d and  or just  which i t o c c u p i e s  11 mm o u t s i d e was  taxon.  down t h e s t i p e .  vertical  plant i t s e l f ,  of a r t i f i c i a l  11 mm  i n order t o  Queen  Starting a t the  a t 70°C f o r 1 week a n d a d r y w e i g h t was  it  locality  were t h e same  vertical  at 1 m i n t e r v a l s  n e r e o c y s t i s p l a n t s i n each  dried  essential  site  formation.  i t s type  and t h e B o l k u s I s l a n d s ,  assessment  were  was  p l a n t s were c o l l e c t e d  P. n e r e o c y s t i s a l o n g samples  at  of spore  a b u n d a n t P o r p h y r a n e r e o c y s t i s was e n c o u n t e r e d ,  Nereocystis  guantitative  Peninsula)  site  was  (13 mm  diameter)  was used  to t i e  17  the  • p l a n t s * t o the  stipes  were  piton ring.  put o u t  i n the  These  field  artificial  i n September  Nereocystis  1974  and  August  from  heavy  1975. Artificial nylon  mesh  cut  pitons,  and  the  m tide  0.4 In  Lam i n a r i a - l i k e into  made  They were a l s o a n c h o r e d  (Table  being  using  approximately  level. to  the  I)  o b s e r v a t i o n s and species  strips.  were  were p l a c e d i n t h e L a j j - n a r i a zone a t  addition  herbaria  plants  field  were  study,  examined  to confirm  s p e c i m e n s i n numerous  to  t h a t I had  supplement  correctly  field  identified  the  studied.  Results  A.,  Poi£hyra  1. A  gardneri  Seasonal brief  Laminaria  to  study the  and  site  stipe  the  by  new  w h i c h was  (Figure 3a). began t o s p l i t as  the  blade  of  the  blade  o f L.  meristematic  Growth distally grew and  was  t o an  transition a  rapid  g r o w t h and  o f Porphyra  late  was  and  i n the  decay of  of the  gardneri.  worn b a c k  November  or  early  heart  formation  a  shape  w i t h i n a month t h e This splitting  At  almost  zone commenced f o r m i n g  characteristic  ( F i g u r e 3b), resulted  *Host*  understanding  setchellii  During  initially  And  annual  occurrence  November.  the  Epiphyte  i s essential  seasonal  December, blade,  Of  consideration  setchellii  establishment the  Occurrence  Made  continued  of the  typical  18  lacerated  b l a d e by F e b r u a r y  During  the f i r s t  relatively  free  ectocarpalean appeared  o r Harch  months  of  of  (Figure 3 c ) .  growth  epiphytes.  Growth  e p i p h y t e s became o b v i o u s  first  on  the  distal  tissue.  became  increasingly  epiphytized  erosion  r e s u l t e d i n the disappearance  first  February  striking, blade to  During  of  early of  the  and  new  was  diatoms  and  summer;  they  the blade  which  autumn  worn of a  blade  away.  the  blade  This  fclade  suitable  substrate  g a r d n e r i appeared  a t t h e end  I2r.Dhy.ra a a r d n e r , i . The  of  by  portions  contained the oldest  for  the  thalli  a t the study being  increase  (Figure  setchellii  margin  soon  produce  the  o f P.  distribution  became  gardneri t h a l l i  densely of  lined  Porphyra  due t o a m a s s i v e  (Figure 4a). colourless  They  patchwork  a month a f t e r submarginal  were  observed  easily  margin  with  that  during  recognized  by  the spermatangia  r e g i o n s became s l i g h t l y  more d e e p l y  production  of  region  carposporangia  their  The f i r s t  spermatangia  vegetative  of  the  gardneri  t h e appearance of the  the c e n t r a l  continued  plant had  a  distal  p r o d u c t i o n and r e l e a s e o f  ( t o be d i s c u s s e d i n P a r t I I I ) . were  very  t h e s p r i n g , and the margins o f  thalli  spermatangia  was  margins o f t h e L a m i n a r i a  l o o s e g e l a t i n o u s appearance along  ( F i g u r e 3d)  monospores  to  throughout  These  characteristic  Their  T h e number  rapidly  1a).  site.  restricted  (Figure 3c).  lainaria  of P o r p h y r a  the  thallus,  thalli  to  late  April  the  fine,  form.  About  the  lateral  pigmented due  to  than the  ( t o be d i s c u s s e d i n P a r t s I I and  III). Thalli  growing  i n the spring  and e a r l y  summer c a n r e a c h  a  19  length  of  20-70 mm on  130  mm  l o n g and  Eisenia  and  a w i d t h o f 60  10-30  arjborea  mm  wide.  generally  much s m a l l e r  The  number  of  L.  setchellii  a p p e a r e d on restricted As  blade  long  as l a t e a s  British site  will  thallus gardneri  (Figures  ( F i g u r e 4c) p l a n t s but  and  in  continue  and  4c,  4d,  other  and  season. as  the thalli being  is  present  (Figure 4b). but  and  sites  Most  I collected  some  to those  4e).  only  study  of  the  cytological  work  present  and  sites.  at  the  observed  S p e c i m e n s from  once  seasonal  reproduction  where i t was  have more r u f f l e d  visited  spermatangium two  grow  September-October,  were s i m i l a r  d i f f e r e n c e s i n the  these  were  instead of  thallus  to  were s i m i l a r i n a l l o t h e r  field  is  i n the  autumn  blade  which  thalli  time s e v e r a l s m a l l  Laminaria  respects. is  occurrence  or  Monterey ( t o be  study  Porphyria  at  the  Botanical  impossible  that they site.  study Beach site these  to  make  combined  with  i n Parts I I , I I I , are the  same t a x o n  However, t h e r e and  formation  in  reproduction.  discussed  g a r d n e r i was  collected  Because  plants,  abundance o f t h a l l i , carpogonium  of  margins than study  i t  I V ) , c o n c l u s i v e l y demonstrated  which  the  At t h i s  occurred  setchellii  found e a r l i e r  Laminaria  at the  comparisons of the The  Laminaria  morphology  tended to  were  culture  old  gone by  Columbia  sites  thalli  are  November.  The JSIfiilXEii  declined  eroded.  as  g a r d n e r i was  largest  they  margins.  Porphyra g a r d n e r i !•  than those  s u r f a c e of the  to the  and  but t y p i c a l l y  L a t e r i n t h e summer t h e  thalli  the  The  Areschoug  were g r o w i n g s u b t i d a l l y .  mm,  in  the  between t h e  were  time  of  plants  at  more a b u n d a n t a t  Point  20  Joe  (type  Also the  locality) thalli  Pescadero (30  mm  north.  at  Point  long  i n November Point  a v e r a g e , up t o 70  (CSUH,  DS,  monosporic,  mm)  than  and  collected  this  being  monosporic  spermatangial,  GMS,  predominance  thalli  and  herbarium that  they  Point  at  farther  California, in  UC, UCSC)  several confirmed  in  whereas  and m o n o s p o r i c .  monosporic  almost  thalli  a t the study  and  the  genus  always  Porphyrella  carpogonium f o r m a t i o n  2.  *Host»  Porphyra observed (Table  Throughout  The  in  site  the  was  and c a r p o s p o r e  on  frequency  most  thalli  examination  Monterey  based  were  J u l y i s i n sharp  An  monosporic.  lateral  populations.  where  spermatangial. in  were  others  i s low compared t o more n o r t h e r n of  1976  had s p e r m a t a n g i a l  specimen i s e x c l u s i v e l y monosporic, a c u r i o u s that  July  by c u l t u r e work done a t t h e  margins,  specimens c o l l e c t e d were  Joe  Some t h a l l i  distal  carposporangial  alqae  as  from  Alaska,  RS, UBC,  confirmed  contrast to the situation were  at  carposporangial  carposporangia  been  well  specimens from  Columbia  GJH,  Hopkins Marine S t a t i o n .  The  as  site.  observations. Most t h a l l i  of  (Figure 4f)  o f herbarium  Oregon, Washington, B r i t i s h  these  Joe  i t was a t t h e s t u d y  a n d M i s s i o n P o i n t were g e n e r a l l y much s m a l l e r  An e x a m i n a t i o n  herbaria  than  area  In f a c t , fact  of  revealed the type  considering  the  details  cf  release.  Species gardneri i s not r e s t r i c t e d on  fifteen  II).  i t s range  one  host,  having  l a m i n a r i a l e a n and one r h o d o p h y c e a n  Saxicolous i t  to  specimens  reaches  optimal  were  not  development  found. on t h e  21  margins of The  Laminaria  •host*  setchellii  species  approximately  +1.4  grow  m to  -3.0  (10-20 mm  long)  fireschoug,  which a r e i n t h e  early the  spring  Porphyra  generally  l a r g e r and  Laminaria  setchellii. all  associated  the  was  the  thalli  o f P.  like  Egregia  by  May  be no  gardneri t h a l l i  species,  gardneri  P.  margins.  Pcstelsia  range  between is  menziesii  thalli  small  {Turner)  range.  found  on  became r e p r o d u c t i v e  host  tidal  species.  Porph y r a g a r d n e r i  g a r d n e r i can  'host'  Laminaria  upper p a r t of t h i s  with the t h a l l u s  trend  the  m.  of E g r e g i a . but  'host'.  On  P.  other  in  'host's'  months  •leaflets'  this  on  cr  In  i n abundance  on  were  on  found  E i s e n i a arbqrea  e a r l i e r than  gardneri One  those  was  on  were on  typically  exception  to  this  p a l m a e f o r m i s R u p r e c h t , where  were g r o w i n g  the  abraded p o r t i o n s  of  the the  stipe.  3.  Artificial  None artificial  4.  of  the  p l a n t was  own  the  Porphyrella  Scagel  Monterey gardneri)  grow P o r p h y r a g a r d n e r i on  the  Distribution  i t i s clear  gardneri)  to  successful.  collections,  literature  Porphyrella  Experiment  attempts  Geographical  distribution.  to  the  host  From my in  Host  herbarium  and  t h a t Porphyra g a r d n e r i  (1957) and to extend  Dawson  from  Peninsula. extended  specimens  the  reports  has  a wide  (1961) r e p o r t e d  i t (as  northern  British  Sparling southern  range  Columbia  (1971, to  as  Hazard  22  Canyon, 120°  San  Luis  53*»).  Setchell (31°  Obispo  S p e c i m e n s on  the  <AHF 77063) f r o m  44»N,  116°  44'H)  County, leaflets  Punta  This  represents  extension  £•  great  gardneri.  water  California cccler  described  coast.  of  water  algae  are  the  Dawson  other  found  Eqregia  and  are  laevigata Mexico  referred  to  population  southern  limit  of t h e (1951)  as w e l l  areas along  typically  there  18'N,  isolated  w i t h i n one  by  Several  (35°  Banda, B a j a C a l i f o r n i a ,  apparently  P u n t a Banda f a l l s  upwelling  of  have been examined  Porphyra g a r d n e r i . a  California  more  of  of  cold  the  Baja  northern,  (Abbott  and  North,  1972). Hynne  (1972)  Porphyrella  gardneri)  Islands,  Alaska.  British  Columbia  into  northern  Murder  Specimens o f  1930, of  (A  Bory  along (52°  in  the  in  the  represent collected  i n the  epiphytic  on  of t h i s  Laminaria  by  Zinova* in  m a r g i n s have b e e n  name t r a n s l i t e r a t e d  on  (1940)  on  to  distributional  of  30  the  Porphyra  reported  the o r i g i n a l  to  Commander  January Academy  been e x a m i n e d .  No  well  7949).  referrable  has  specimen).  UBC  the  western  cf  Islands  E. K a r d a k o v a on  Leningrad  Aleutian  range  by  Botanical Institute  U.S.S.H. i n  photograph  its  the  11»£,  are  (as  University  173°  Island  58»E)  the  Aleutian  48»N,  Bering  165°  and  deposited of  from  12 * N,  specimen  Science  Author*s  west  Island  56455 i s a  1  Island  Porphyra n e r e o c y s t i s reported  (55°  and  and  Attn  Porphyra g a r d n e r i limit.  Amchitka  P h y c c l o g i c a l Herbarium extend  longipes  Islands  from  Por.phy.ra g a r d n e r i  C o l l e c t i o n s deposited  Alaska  Point,  Laminaria  reported  as  species  f o r the  Sinova.  UBC  Sea  23  of  Okhotsk  1964), Oh mi  (Zinova,  or  the  (1963) do  Laminaria, confirmed Strait  not  »  M.  P.  gardneri  1941).  (Vozzhinskaya, and  Japanese s p e c i e s of Porphyra  on  Kurogi  British  Island  (1960)  (personal  Porphyra g a r d n e r i  is  communication)  also  Columbia. from  Tokida  absent  A list  throughout  of  has  frco  the  representative  this  geographical  i n Appendix I I .  Porphyra n e r e o c y s t i s  Seasonal  Occurrence Of  Nereocystis  luetkeana  s u r v i v e f o r as  study  site  at  essentially growth.  the  Buprecht  was  and  pneumatocyst had  reduced.  as  18  The  of  during  on  Setchell  species during  the  the  appeared  period  and  showed  at  the were  of r a p i d  stipe  Membranippra pneumatocyst.  fenestrata Postels et appeared  By J u l y most o f t h e and  plants  plants  bryozoan  laminae  surface  stipes  a few  The  Enteromorpha  August. the  although  March.  e t G a r d n e r , U.  of  'Host*  It first  J u l y , growth o f t h e  noted  reached  And  annual,  months.  beginning  In J u n e and  stenoehyl,la.  plants  long  Epiphyte  i s an  epiphyte-free  megfaranacea Diva  (Nagai,  m e n t i o n any  of Georgia,  1.  can  Sakhalin  Islands  Dr.  this.  range i s given  E  Kuril  and  specimens of  1954) ,  stipe  on  the  Nereocystis  growth  was  s i g n s of a b r a s i o n , but  much  were  not  epiphytized. Once N e r e o c y s t i s phase, e s t a b l i s h m e n t to  early  October  ectocarpalean  reached  the  of epiphytes a  bacterial  epiphytes  became  end  of  was  rapid.  and  diatom  established  its By  stipe late  growth  September  'scum* f o l l o w e d on  the  by  stipe.  24  Throughout  the  progressively of  the  stipe  almost winter  autumn  A few  plants,  were e a s i l y  study  the a t low  narrow  site  recognized  5a}.,  folds.  Large,  portion  tide.  The  The  During colourless  90  their  the  dense c o a t i n g  mm.  thalli  which  up  which  have  a  slowly  t o 2.7  t o the  the  lateral  and  floats  without  m long  production  spermatangial  continued  of  to a  the  the  of  carposporangia submarginal  v a r i e d i n width from  the  and  distal  region  carposporangia  marginal  never observed  Humford  began of  or  forming  Conway  early in  the t h a l l u s  strips  in small  (1973a) and  January  areas  or  became  (Figure  i n a basal d i r e c t i o n ,  were j o i n e d i n p l a c e s t o t h e  end  point  were  thalli  of  February, the  ( t o be  (Figure  10  mm the  5c).  formed,  (Figure  microscopic et  5b).  leaving a  narrow, s u b m a r g i n a l s t r e a k s o f s p e r m a t a n g i a a l s o  r e p o r t e d by  the  wrinkles  Long,  were  It  cn  ( F i g u r e 1b)  spermatangia  strips  to erode  vegetative c e l l s  Spermatangia  on  characteristic  tapers  margins of  They b e g a n t o e r o d e i n and  stipe  t i p of  as  of  February.  January, due  of the  shape  appeared  distal  and  winter  November t o e a r l y December.  young  mature t h a l l i  marginal  thallus  by  margins are s t r a i g h t ,  i n J a n u a r y and  These  survived  Porphyra n e r e o c y s t i s f i r s t  upper  ribbon  (Figure  found  which  -  the  on  thereby  erosion  the  Nereocystis stipes in late  long,  to  p l a n t s and  Breakdown and  became  complete e l i m i n a t i o n o f the N e r e o c y s t i s beds d u r i n g spring.  the  plants  the  At  surface  the  c o n t r i b u t e d to  and  weakened  epiphytes.  grew  winter  more h e a v i l y e p i p h y t i z e d .  storms i n t a c t ,  the  and  5d).  patches  a l . (1975).  By  carpogonia  and  distal discussed  and  lateral  i n Part I I ) .  25  In  elder t h a l l i  mottled  the c a r p o s p o r a n g i a  appearance  in  spring  and  the f i e l d , early  the  number  stipe  These <C.  Agardh) C.  £•  (OBC  Porphyra approximately New  Agardh  thalli  (OBC  two  collected  57194),  October  (J.  in  Woessner,  collected (OBC  in  (OBC  with  a  observed  on  gt  nereocystis.  55942),  P.  miniata  p e r f o r a t a J . Agardh Abbott  {OBC  57220),  (OBC and  57195).  young  thalli  the  South  Friday  Porphyra than  Strait  September Pender  Georgia  did  communication,  at  at  August  (OBC  Juan I s l a n d ,  also  the  site.  Sidney,  Island  H a r b o r , San  nereocystis  i t  of  i t d i d a t the study  e a r l y as  and  at  California personal  than  as  1434)  in  appeared  study 1S77).  i n November a t t h e M o n t e r e y  site I  Peninsula  57196). At  the other  sites  along  the  Columbia the appearance of t h a l l i at  Porphyra  (UBC  57219), P.  months e a r l i e r  (OBC  earlier  with  n e r e o c y s t i s appeared  Vancouver I s l a n d  Washington  or  57201) .  were  and  July  associated  s p e c i e s were a l s o  along  Hollenberg  to  year.  Porpfryra g a r d n e r i  smithii  thuretii  a  i t i n abundance i n t h e  always  the p r e v i o u s  Nereocystis  November  collected  of other Porphyra  of  £•  from  summer when i t i s  included:  57222),  lasted  I have  N e r e o c y s t i s p l a n t from A  tips  (Figure 5e).  P01jp.h7.ra n e r e o c y s t i s August  o f t e n gave t h e d i s t a l  the  time  occurrence  of c o l l e c t i o n  similar  to the  and  suggests  study  site  outer their  coast  of  reproductive status  t h a t t h e y have a plants.  British  seasonal  26  2.  Vertical Distribution  Porphyra  nereocystis  depending  on  plants  concentrated  are  largest At t h e  its  varies  position on  plants t y p i c a l l y l o w e r end  (10-100 mm)  of  and  thalli  *Host»  the  P.  with  57197).  the  4.  can  Artificial  The  best  are  Host  material  surgical  rubber.  1975  l a s t e d 14  months.  some  fiollenbergia Postels  et  of  thalli  They had  the  was  of  stipe,  first  1-2  the  curious  t o one  m  Host  with  the  {Table I I I ) .  thalli  are  feature  i s that  s i d e of the  f o u n d e x c l u s i v e l y on o f a few  blades  there are  small on  stipe.  reports  the  found  s t i p e of  growing  integrifclia  i n the  tested  Sory  literature  The  for imitating a Nereocystis 'plant* I  I t supported  Huprecht  and  typical this  a  the  i n the  that .  a  Alaria  making  in of  margjnata  var.  small  artificial stipe  s h a p e o f young P.  field  stenophylla.  ligulata few  stipe  growth  Jilya  Sollaston,  I n November*  stage,  out  magnificent  Desmarestia  on  put  included:  (Harvey)  appeared  on  Experiment  subulata  them grew b e y o n d  thalli  of Macrocystis  which  ( L i g h t f o c t ) lamouroux. long)  limited  abundance  grow s a x i c o l o u s l y , I n e v e r o b s e r v e d t h i s  was  algae,  i n the  another  and  Nereocystisstipe.  upper 3 m  exception  Although  nereocystis  the  size  Species  heavily epiphytized  (UBC  its  distribution  Porphyra n e r e o c y s t i s Nerepsystis  the  vegetative,  most s t i p e s t h e  3.  on  being  the  in  ligulata  (30-40  mm  (UBC  57221).  n e r e p c v s t i s . but  none of  positive  identification  27  impossible.  5.  Geographical  From  my  own  herbarium  Distribution  collections,  specimens,  the  known  Forphyra  nereocystis i s : Staraya  Islands,  Alaska  to  Box  (53°  r e p o r t s i n the  37.4»N,  Bay,  165°  and  <35° 10«M,  south  120°  occasionally  45*8)  1976).  Zinova  (1940)  San  in  l c n g i p e s and  Commander i s l a n d s ,  (59°  1961).  Drift  165°  a b l e t o examine t h e s e  specimens,  which  are  Botanical  of  from  personal  on  re-investigating Nereocystis copses  plant).  communication)  drift  i n the  that  I  feel  attemped  or impossible to obtain  in  the  Laminaria  the  specimen  (UBC  57206, i s  ( P . A.  Lebednik  this  report  was  that t h i s  zone  Island,  o f U.S.S.B. i n  and  , of  unknowingly  would be  (1940) s p e c i f i c a l l y  sub-littoral  on  I have been  deposited  Commander I s l a n d s  because Zinova  have  and  s p e c i m e n from  Some workers  specimens.  grows i n t h e  I  (Abbott  58*E) .  nereocystis  feel  which make i t d i f f i c u l t  through."  are  Porphyra g a r d n e r i : however,  of t h i s  Nereocystis luetkeana based  the  N e r e o c y s t i s i s r e f e r a b l e t o P.  a colour slide  California  at Bering  t h e Academy o f S c i e n c e  I indicated earlier,  lcngipes i s actually  UBC  t h a t P o r p h y r a n e r e o c y s t i s grows  12«N,  As  22'H,  specimens  California  (55°  Leningrad.  of  northeast  147°  County,  Nereocystis luetkeana  U.S.S.E.  Institute  26776),  58'N,  Obispo  southern  reported  distribution  30.6«W, UBC  Luis  (Dawson,  found  Hollenberg,  Laminaria  to  and  Onalaska I s l a n d , A l e u t i a n  P o i n t , Montague I s l a n d , A l a s k a  25836)  literature  worth  stated that  forms  "thick  f o r a boat t o  pass  further information  on  28  this  from  C.  point,  but without  A list  of representative  throughout  Por£hy_ra  1.  Occurrence  This l i t t l e  known a l g a  in  February  along  with Porphyra  1975  species  several  Porphyra abundant after  May.  ruffled  I  have  thalli  was  give  spermatangia  P.  distroroatic  short  identified  by  collected  this  first  streaks  i n the l a t e r a l  not  is  readily  not  encountered  the  strongly  distinguishes  Superficially  a resemblance  t o P.  the  miciata;  on t h e b a s i s o f  spermatangia  do s o u n t i l  noticeable  in  but i t was  their  constructions, respectively.  producing  increased  p a t c h e s and  feature  nereocystis.  m a r g i n a l band a t t h e t i p o f t h e gradually  luetkeana from  I t was  which  P. t h u r e t i i  most d i d not are  study  plant  c o l l e c t e d i n January,  c a n r e a d i l y be s e p a r a t e d  and  at the  fertile  was  or early A p r i l .  from  thalli but  one  subsequently  (Figure 6a),  margins  encountered  52019)  margin  monostromatic  apex and  The  most c h a r a c t e r i s t i c  A few  i n Appendix I I .  occasions.  March  however, t h e two  January,  first  The  vegetative ruffled  (UBC  thuretii  until  was  nereocystis.  Mumford, J r .  on  i s given  nereocystis  on t h e s t i p e o f N e r e o c y s t i s  collection T. F .  range  of Porphyra  thuretii  Seasonal  this  specimens  i t s geographical  site  Dr.  success.  length  observed  F e b r u a r y or March.  as  thallus  were  a  thin,  of spermatangia  submarginal areas  and formed  on  The  colourless  (Figure 6b).  basally,  in  This  older  band  thalli  at the t h a l l u s  ( F i g u r e s 6c  and  6d).  29  The  specific  formation  time  i s not  J a n u a r y , but  cf  the  known.  initial  Fertile  stages  thalli  of  were  most d i d n o t f o r m c a r p o s p o r a n g i a  carpogonium collected  in  u n t i l February  or  March. Porphyra 150-220 mm by P.  105  thuretii  long  mm  and  did  occur.  less  larger  pronounced  Barkley  Sound  the  50-80 mm  t h u r e t i i from t h e  generally  in  Herbarium  t o 770  marginal  been  for their  larger size.  typical  of the  (Figure  6e).  2.  i n the  thallus  one  case  (1944) and  was  found  Smith  p h a e o p h y c e a n and Barkley  Sound  epiphytically contrast,  occurs on  (1944). 3  mm  year  may  of to  wide) and  be  be  mm  GMS)  noted  be  have  specimens  from  that  the  (May-June)  and  an  explanation  i n A p r i l were  I have o b s e r v e d  rock,  on  region  I t has  only  cn  P.  more  in Barkley  Hhodophyceae.  the  host  Sound  algae  of  the  on  (Table was  species,  reports of  been o b s e r v e d  thuretii  members the  7 different  confirming  rhodophycean  specimens from  members o f t h e  so t h i s  390  Species  Porphyra t h u r e t i i in  i n the  to  were f o u n d  the  I t should  later  drift,  than  Specimens c o l l e c t e d  size of  *Host'  6d).  230  up  (DC,  region  l o n g and  ruffling  ( F i g u r e s 2c and  floating  mm  is typically  thalli  specimens  Monterey P e n i n s u l a  (up  Sound a r e a  wide, although  M o n t e r e y p l a n t s were c o l l e c t e d had  Barkley  and  Dawson  1 seagrass, IV).  observed  In  3  the  growing  Laminariales.  In  M o n t e r e y r e g i o n were e p i p h y t i c on  30  3.  Artificial  One  •Host* E x p e r i m e n t  p l a n t o f Porehyra- t h u r e t i i  on  the  and  reached reproductive  4.  artificial  Nereocystis  Geographical  (UBC  57199) which  p l a n t grew t o 146  Bennell  132° 30.7*H,  Island  sites  Barkley  in  1 2 5 ° 11. V B , Doty  OBC  Sound,  reported  (36° 30* N,  (1952)  Pacific  in  t h e h e r b a r i a o f UC  once  (53° 22.65*N,  Kyuguot  Channel,  54446) and  Island  125°  collected  (48°  14.4*B, OBC  two  50.1»N,  57200) .  a t C h e t c o Cove i n Oregon  a different and  representative i n Appendix I I .  i s known o n l y  (Dawson, UC  1944),  95596)  from  the  and  Santa  Monterey  Carmel  Bay  791973).  California,  and s p e r m a t a n g i u m  given  i t  121° 57*H,  reported  in  A  Vancouver  122° -00*9)  (36° 33*N, 1 2 1 ° 5 6 « 1 , OC  represent  Islands  Cove,  Porphyra t h u r e t i i  (36° 58*N,  Baja  been  18»W).  In C a l i f o r n i a ,  Dawson  Volcanic  has  (49« 58.7'N, 127° 13.9»W, UBC  (1947)  Peninsula  Charlotte  52019; 48° 49.8»N,  (42° 00*N, 124°  Cruz  Queen  55302),  Vancouver  i n length  Distribution  Sound, UBC  mm  maturity.  In B r i t i s h Columbia, Porphyra t h u r e t i i at  appeared  and  P.  thuretii  Mexico.  AHF,  from s e v e r a l l o c a t i o n s  These s p e c i m e n s ,  were e x a m i n e d  taxon because o f t h e i r carposporangium specimen  list  deposited  and i n my small size,  opinion shape,  distribution. of Porphyra t h u r e t i i i s  31  Discussion  A.  Porphyra  gardneri  Although months  of  the  species. of  the  Its  been f o u n d  year,  at t h e s t u d y  Porphyra  monospore c y c l e  gardneri results  p o p u l a t i o n t o some e x t e n t  persist the  i t has  l o n g e r than  monospore  foliose  phase  to  in  from  has  year-round  s p e r m a t i u m and studies  noted  spore  formation  of Porphyra  Dixon,  1968;  Nichols,  in  to  photoperiod  35°N  have been o b t a i n e d (List, daily  1966),  (National  36°  30 * N  the  21st  and  48°  50.1»N  from  and  day  Bangia 1969;  i s the  i s used  Council,  (Monterey P e n i n s u l a , t y p e  the  and and  Therefore  result  of  In Table  in i t  i n the times  o f e a c h month i s  cf the  V  the  compared  Data f o r 35°N  Meteorological  Tables  have been t a k e n  Astrophysics  because i t i s  Culture  i s important  latitude.  data  of  Sommerfeld  (study s i t e ) .  study-site  Fcrther  (Richardscn  variation  the Smithsonian  the  freguency  latitude.  sunset t a b l e s f o r V i c t o r i a ,  Eesearch  L a t i t u d e 35°N  with  of  suggestion.  formation.  that  enable  area.  photoperiod  carpospore  on  whereas  s u n r i s e and  and  that  variation  latitude  this  to  predominance  this  v a r i e s with  suggest  re-establishment  The  time  carpogonium f o r m a t i o n  duration of daylight for  the  Bichardson,  shown  monospore and  s p e r m a t a n g i u m and annual  and  have  reasonable  that  ( I w a s a k i , 1961)  Dixon  1973)  controlling seems  been  10  spring-summer  M o n t e r e y may  seasonal studies are reguired to confirm It  a  i n the  species.  plants  exist  is  throughout  i n t h e autumn, e n a b l i n g i t  most P o r p h y r a  cycle  site  British Branch,  closest  from  Columbia 1974).  latitude  l o c a l i t y ) f o r which d a t a  to are  32  given.  There  is  approximately  duration of daylight June,  but  present  this  between t h e s e two l a t i t u d e s  i s  accurate  the study s i t e  daylength  was  carposporangia, longer  spermatangia  approximately formed  daylength  duration  was  of daylight  strongly freguency  Whether  temperature  The vertical which  be  21st  of the  may  also in  position  may  exposure,  be  be s e e n  in a  reached  Part  a  (1961)  slightly  be  Table  between V,  this  a t 3 5°N on 2 1 s t J u n e .  III.  of  Higher  seawater and N i c h o l s  temperature  favoured  carpospores,  size i s  plants.  a photosynthetic  Sommerfeld  higher  thallus  result  Kurogi  to  from  factor.  on t h e s h o r e  the  when a  the then  f o r m a t i o n i n the Monterey  Bangia  in  this  estimated  o f monospores r a t h e r t h a n variation  Carpogonia,  photoperiodic or just  be  when  photoperiod i s responsible f o r the  discussed  found t h a t  production  As w i l l  suggests that  appeared  14 h r .  i s only just  i s a truly  will  first  reached,  of carpogonium  this  effect  (1973)  the  f o r t h e purposes  a s h o r t time a f t e r  14 h r 30 min and 15 h r .  low  enough  on  i n the  discussion.  At  This  a 10 m i n u t e d i f f e r e n c e  o f Porphyra  an  g a r d n e r i with  interesting  phenomenon  growth i n h i b i t i o n by p r o l o n g e d  noted  that  growth  of  Porphyra  y e z o e n s i s was more r a p i d a t l o w e r i n t e r t i d a l l e v e l s . As  a  result  c o n s i d e r a b l e number in  o f my s t u d y ,  o f •host* s p e c i e s ,  the L a m i n a r i a l e s .  specific The  Whether t h e r e  compound p r o d u c e d striking  P. g a r d n e r i i s now known  from  a  a l l b u t one o f which a r e  is  a  requirement  for  a  by t h e L a m i n a r i a l e s i s n o t known.  marginal d i s t r i b u t i o n  blades of Laminaria s e t c h e l l i i  i s probably  o f P. g a r d n e r i on t h e the  result  of  the  33  tearing spore  of the  germination  appears P.  Laminaria  to  be  and  In  case  for  growing  recent  Harveyella that  spore  wound a r e a on  the  the  on  work  mirabilis  producing  a wound s i t e  attachment can  been wounding c a u s e d  her  found  rhizoid  the  g a r d n e r i i s found  t h e r e has  blade  of  plant  p l a n t s , where  a b r a s i o n o r some o t h e r  means.  red  Schmitz  caused  algal  e t Beinke,  e s t a b l i s h m e n t depended host  also  where  the  the  This  thallus  on  (Beinsch)  e t h e r 'host*  areas by  occur.  i n which  by  on  parasite  Goff  . (1975)  the  presence  of a  grazing  isopods  and  amphipods. The due  absence o f e p i p h y t e s  t o r a p i d growth  production  of  1966;  The intriguing unigue  the  and  S i e b u r t h and  one.  position  which  some  (Craigie  phenomenon  of  Harlin  material  or  (e.g. p h e n o l i c s )  are  1964;  HcLachlan  and  host s p e c i f i c i t y  plant occupies i n the  epiphyte  The  found  rock  would n o t  host.  be  (1973a) d e m o n s t r a t e d t h a t  obligate  t h a t i t may  may  fact  and  1969)  epiphytism  reguires.  suggests  which  Jensen,  t h a t the h o s t  and  of  blades  surface  Mclachlan,  Hollenberg on  young L a m i n a r i a  s l o u g h i n g o f f of  exudates;  known t o be t o x i c Craigie,  and  on  Smithora  that Porphyra grow on  i t  was  i s an the  environment  naiadum gardneri  the a r t i f i c a l  have a p h y s i o l o g i c a l r e g u i r e m e n t  (Anderson) was  not  Laminaria for  the  34  E.  Porphyra  My winter  field  it  and  The  summer  Hollenberg  reports  i s that  i s not  ether  because  *burnt  winter  can  persist  is  not  are  off*  and  Porphyra  c a r r i e d away.  round i t i s easy to  and  nereocystis spring  a different  it  et  occurs  (1977)  that  phenomena.  The  is a subtidal  plant  as  (Mumford, 1 9 7 5 ) . that  Without c o l l e c t i n g  get  Cole  months  summer p r o v i d e d  Smith  Woessner  (1975) t h a t  o f Conway and  the  Porphyra s p e c i e s the  reports of  a t t r i b u t a b l e t o two  during  well info  nereocystis i s a  the  (1976)  o f Conway e t a l .  through winter  i s a summer s p e c i e s  first it  T h i s i s i n agreement with  Abbott  (1977).  from  s t u d i e s i n d i c a t e that Porphyra  species.  (1944), al.  nereocystis  happens  As its  a result i t host  plant  at a s i n g l e s i t e  concept  cf  to  the  year  seasonal  occurrence. The  second  much e a r l i e r  appearance  Georgia  and  is  autumn  an  thought I  phenomenon  vicinity  Just  what  responsible P.  Most the  workers  release.  stabilizing  effect  Parsons of  coast.  this  In  the  of  Strait i t  i t this early I  of Georgia  they a r e ( t o be  et a l .  however,  has  region  probably  1972)  of  involved in  Part  have i n d i c a t e d  factor required  noted  input  are  occurrence  discussed  is a critical  freshwater  the  Strait  species;  seasonal  (1965)  is  conclusion.  Strait  the  Bird  the  encountered  c l e a r , but  (e.g.  confusion  in  different  i n the  seawater temperature  ccnchcspcre the  not  a  supported  altering  are  outer  t r i g g e r i n g conchospore r e l e a s e  III). that  not  conditions  for  nereocystis  with  has  the  with  caused  nereocystis  When I f i r s t  dealing  subseguent study  P.  t h a n on  plant.  was  of  w h i c h has  t h a t , because from  the  for of  Fraser  35  River, in  the s p r i n g phytoplankton  the  Strait  factors  involved  Nereocystis year  of G e o r g i a  and  t h a n on  here are  p l a n t s i n the begin  to  low  Strait  senesce  day  t i d e s i n the  of  the  i n the  the  cuter coast  Strait  in  higher  floating  on  the  earlier  Strait  t h a n i t does on  observed that  i t growing  such  found al.  to  t o be  P.  (1975,  opposed  at the  P.  and  low  are  F i g . 19a, miniata.  as  being  reports of  P.  n e r e o c y s t i s on Kylin  nereocystis.  other  Porphyra  fly species  agree  the  in  studies are  plants Strait.  reproductive  being  I have  Mumford  the  UBC  never  (1973a)  P.  pools  were  Conway  nereocjjstis  of  P.  gardneri  et are and  has  been  responsible  for  other  than  Nereocystis.  For  P.  f o r a young o r  gardneri  on  dwarf f o r m  of  have a l s o shown t h a t a number  capable  of  herbarium  f i g u r e d by  as  mistook  Ruprecht  on  This  misidentifications  33206)  (1941)  morning  specific,  with  young t h a l l i  hosts  middle  for  in mid-intertidal tide  nereocystis  Pterygophora c a l i f o r n i c a P.  »host*  Misidentification  miniata  example,  in  is  coast.  specimens  UBC  coast  1975).  a l s o becomes  probably  The  the  plants.  i n the  insolation tide  Strait  and  saxicclous  P.  cuter  1969b; Humford,  outer  rocks  miniata.  r e f e r r a b l e t o P.  the  in  coast  t o e a r l y i n the  Several  be  earlier  to Nereocystis luetkeana.  reports  fofihXEa l i f i i a t a . reported  on  The  as  the  essentially  nereocystis.  summer o c c u r  Porphyra n e r e o c y s t i s i s remarkably restricted  the  than outer and  earlier  Perhaps  s p r i n g and  surface in  weeks  coast.  appear  earlier  desiccation  nereocystis  outer  also  ( Krishnamurthy,  results  ESlEhlE*  the  4-6  a l s o a f f e c t i n g P.  A n o t h e r d i f f e r e n c e between t h e that the  bloom o c c u r s  of  g r o w i n g on  the  stipe  of of  36  Nereocystis  and  could  potentially  be  mistaken  for  P. n e r e o c y s t i s . Porphyra n e r e o c y s t j s had  terminated  encountered by  a  i t s rapid stipe  until  heavy  was f o u n d o n l y  t h e autumn  layer  with  P. g a r d n e r i .  by  abrasion  on  growth.  and w i n t e r  Nereocystis  Such p l a n t s were n o t  and were  characterized  o f d i a t o m and e c t o c a r p a l e a n  i t is felt  or  that  grazing  a wound  is  which  epiphytes.  on t h e s t i p e ,  important  for  As  caused  conchospore  establishment. The one  reason  side  of  f o r P. n e r e o c y s t i s u s u a l l y b e i n g the  stipe  observed a t low t i d e . stipe is  is  floating  exposed  to  desiccation, !!•  becomes o b v i o u s when t h e k e l p  At t h i s t i m e ,  at the surface.  the which  restricted  air  the  the  The h a l f o f t h e s t i p e  that  would  tend  to  a  3-5  great  inhibit  m  bed i s of  experiences  upper  to  amount  the  growth  of of  S§£§2£2§tis. The  epiphytic habit of  advantages. free  from  submerged  The  plant  grazing,  and  Pot£byr§ n e r e o c y s t j s  i s on a s u b s t r a t e which  but near t h e s u r f a c e  keeps  has  obvious  which i s r e l a t i v e l y  the  plant  constantly  and a d e q u a t e i l l u m i n a t i o n .  37  C.  forghyra  My  thuretii  field  observations  winter-early (1944) and At  spring  Dawson  present  rather  patchy;  case.  This  grows  in rather  in £•  the  nereocystis, The  fact  species,  as  the  however, has  I  do  It P.  is  also  miniata  that  Porphyra  It P.  host  differences  at  appears  think t h i s i s actually  the  overlooked  because  further  be the  between  but  the  to  and  make  Peninsula  these p l a n t s  and  a  early  i t  for  by  the  a  the  This  that i t  was  Nereocystis.  seasonal  i n order  study s i t e  thallus size,  physical  offers. fact  and  artificial  physiological  host  Phyllospadix  Monterey  marginal r u f f l i n g  have  the  just r e q u i r e s  which  desirable  mistake  grew cn  not  substantiated as  to  it  perforata.  does  environment  h o s t s as d i v e r s e  thuretii  degree of  the  the  would  Smith  be  thuretii  plant,  on  by  is a  to  been  o r P.  on  found  reported  thuretii  possible  dependance  is  been  thuretii  e x p o s e d a r e a s where c o l l e c t o r s s e l d o m go  i t  conclusion  not  probably  suggests that  in  has  d i s t r i b u t i o n o f P.  Nereocystis  position  Porphyra  (1944).  species  year.  i n d i c a t e that  study  to see  plants,  are  of  i f the in  the  significant.  38  Table  AHF: CSOH: ES: GJH: GMS: PS: UBC: UC: UCSC:  I.  Herbaria  from w h i c h s p e c i m e n s  were  examined  A l l a n Hancock F o u n d a t i o n , U n i v e r s i t y o f S o u t h e r n California C a l i f o r n i a S t a t e U n i v e r s i t y a t Humboldt D u d l e y H e r b a r i u m , S t a n f o r d ( s p e c i m e n s now i n UC) P r i v a t e H e r b a r i u m o f G e o r g e J. H o l l e n b e r g G i l b e r t Morgan S m i t h H e r b a r i u m ( H o p k i n s M a r i n e S t a t i o n ) P r i v a t e H e r b a r i u m o f R o b e r t B. S e t z e r U n i v e r s i t y of B r i t i s h Columbia U n i v e r s i t y of C a l i f o r n i a at B e r k e l e y U n i v e r s i t y of C a l i f o r n i a a t Santa Cruz  Table  II.  *Host»  p l a n t s of Porphyra  Host  Acc,L  Phaeophyceae A l a r i a marginata P o s t e l s et Ruprecht Egregia laevigata Setchell E g r e g i a m e n z i e s i i (Turner) Areschoug E i s e n i a arborea Areschoug fljgdophyllutt sessile (C. Agardh) S e t c h e l l Laminaria qroenlandica Eosenvenqe L. s e t c h e l l i i S i l v a i* s i n c l a i r i i (Harvey) farlow L* y e z o e n s i s Mi ya be Lessonioesis l i t t o r a l i s (Farlow e t S e t c h e l l J Reinke Macrocystis pyrifera ( L i n n a e u s ) C. A q a r d h Nereocystis luetkeana (Uertens) P o s t e l s e t Ruprecht IiSU£op_hycus g a r d n e r i S e t c h e l l e t Saunders P o s t e l s i a palmaef q r m i s Ruprecht Fteryggphqra c a l i f p r n i c a Ruprecht Rhodophyceae Schizymenia  pacifica  Kylin  MQJL  gardneri  Location  on h^ost  OBC 54894  midrib  AHF 77063 OBC 54889  tips of leaflets tips of leaflets  OBC OBC  blade blade  54890 54893  margin margin  OBC 54129  blade margin  OBC 54 887 OBC 49214  blade margin blade margin  OBC 54202 UBC 54 883  blade margin blade margin  GMS  13042  blade  GMS OBC  103 55942  b l a d e margin stipe  UBC  54886  torn blade  UBC  55751  stipe  UBC  54885  sporophyll  margin  A b b o t t 2516 t h a l l u s i n GMS  margin  margin  margin  40  Table I I I . per  Igter 1 2 3 4  (top)  Biomass (grams d r y w e i g h t ) o f P o r p h y r a v e r t i c a l meter o f n e r e o c y s t i s s t i p e .  study  site  98 46 18 negligible ( t o t a l of 3 Nereocystis plants)  Queen C h a r l o t t e  nereocystis  Islanas  43 67 11 negligible (total of 4 Nereocystis plants)  41  Table IV.  Host  p l a n t s of Porphyra  Host A r t i f i c i a l host Saxicolous Sea g r a s s e s fhyllosEaaix  sp.  Phaeophyceae Sgjcegia m e n z i e s i i ( T u r n e r ) Areschoug E i s e n i a arborea Areschoug Nereocystis luetkeana (Mertens) P o s t e l s e t Euprecht Bhcdophyceae Gastroclonium c o u l t e r i (Harvey) Kylin Gracilaria sloestedtii Kylin J i c r o c l a d i a c o u l t e r i Harvey (tentative i d e n t i f i c a t i o n from f r a g m e n t o f h o s t )  thuretii  A c c . no.  L o c a t i o n on  UBC 57199 OBC 55302  upper upper  UBC  on t h e l e a f  54447  host  stipe subtidal  UBC 57198  on t i p s o f l e a f l e t s  UBC 57202 DEC 57 200  blade margins stipe  GMS  general  1410  UC 791970  general  UC 95607  general  table  V.  A n n u a l v a r i a t i o n i n d u r a t i o n o f d a y l i g h t ( h r . and min.) on t h e 2 1 s t d a y o f t h e month a t 35°N and 4 8 ° 50.1'N ( s t u d y s i t e ) .  35°N iMx liSjl January February March April May June July August September October November December  10, 11, 12, 13, 14, 14, 14, 13, 12, 11, 10, s.  11 08 09 16 09 31 10 18 12 07 11 48  Study s i t e lira. jilbJ. 8, 58 10, 32 12, 12 14, 00 15, 28 16, 07 15, 32 14, 04 12, 17 10, 32 8, 59 8, 17  43  Figure  1a.  M o n o s p o r i c t h a l l i o f P o r p h y r a g a r d n e r i g r o w i n g on t h e b l a d e m a r g i n s o f Lag j. p a r i a s e t c h e l l i i . The t y p e s p e c i m e n (DS 306401 i n UC).  Figure  1b.  A 2.7 m, f e r t i l e s p e c i m e n o f Pgrp_hjrr_a nereo.cxs.ii§ c o l l e c t e d a t t h e s t u d y s i t e (UBC 5 7 1 8 0 ) .  Figure  1c.  The  l e c t o t y p e of Porphyra t h u r e t i i  (UC  791973).  44  45  F i g u r e 2a.  The f i e l d work was c a r r i e d o u t i n B a r k l e y Sound on t h e west c o a s t of V a n c o u v e r I s l a n d ( i n s e t ) . The s t u d y s i t e was l o c a t e d o f f D i a n a I s l a n d ( c i r c l e d ) , n e a r t h e B a o f i e l d M a r i n e S t a t i o n (from C a n a d i a n H y d r o g r a p h i c S e r v i c e C h a r t #L-3001),  Figure  A e r i a l p h o t o g r a p h o f D i a n a I s l a n d and t h e s t u d y site. The d i r e c t i o n of t h e open P a c i f i c Ocean i s i n d i c a t e d by t h e arrow, (from B. C. G o v e r n m e n t a i r p h o t o g r a p h #BC 7238-182).  2b.  46  47  Figure  3a.  C h a r a c t e r i s t i c h e a r t - s h a p e d new b l a d e g r o w t h o f L a m i n a r i a s e t c h e l l i i (OBC 54895) .  Figure  3b.  E a r l y s t a g e i n the s p l i t t i n g L. s e t c h e l l i i (OBC 5 7 2 0 4 ) .  Figure  3c.  B a t u r e , l a c e r a t e d b l a d e o f L. s e t c h e l l i i w i t h t h a l l i of Porphyra g a r d n e r i along i t s margins (OBC 5 4 8 9 6 ) .  Figure  3d.  Honosporic  thalli  of the blade of  o f P. g a r d n e r i  (UBC  54819).  49  Porphyra  gardneri  Figure  4a.  T h a l l u s o f P o r p h y r a g a r d n e r i w h i c h has s p e r m a t a n g i a , c a r p o g o n i a and young c a r p o s p o r a n g i a along i t s l a t e r a l margins. Monospores a r e s t i l l b e i n g r e l e a s e d a l o n g the d i s t a l margin (OBC 54821}.  Figure  4b.  O l d , l a c e r a t e d , and e p i p h y t i z e d t h a l l u s o f L a m i n a r i a s e t c h e l l i i w i t h a few t h a l l i of P o r p h y r a g a r d n e r i on i t s m a r g i n s ( a r r o w s ) . The new L a m i n a r i a b l a d e i s j u s t b e g i n n i n g t c grow (OBC 5 7 2 0 5 ) .  F i g u r e s 4c-e. Morphological v a r i a t i o n i n t h a l l i of P. g a r d n e r i which a r e s p e r m a t a n g i a l and carposporangial. Note t h e c h a r a c t e r i s t i c p a t t e r n f o r m e d by t h e s p e r m a t a n g i a a l o n g t h e l a t e r a l margins. F i g u r e 4 c : OBC 54138, from B o t a n i c a l Beach; F i g u r e s 4d and 4e: OBC 54891, f r o m C a p e Beale. Figure  Scale  4f.  bar  T h a l l u s o f P. g a r d n e r i from t h e t y p e l o c a l i t y . T h i s p l a n t i s r e l e a s i n g monospores a l o n g i t s d i s t a l m a r g i n and s p e r m a t i a and c a r p o s p o r e s along i t s l a t e r a l m a r g i n s (OBC 54818).  =  10  mm  on  Figures  4a,  4c,  4d,  4e,  4f  51  Porphyra  nereocystis  Figure  5a.  Young v e g e t a t i v e t h a l l i a r e c h a r a c t e r i s t i c a l l y l o n g and s t r a p s h a p e d , t a p e r i n g g r a d u a l l y t o a p o i n t (UBC 57187) .  Figure  5b.  The f o r m a t i o n o f s p e r m a t a n g i a b e g i n s a t t h e l a t e r a l m a r g i n s and p r o c e e d s b a s a l l y . These r e g i o n s o f t h e t h a l l u s a r e e a s i l y r e c o g n i z e d by t h e i r p a l e y e l l o w t o w h i t e c o l o u r (OBC 57188).  Figure  5c.  as a r e s u l t o f s p e r m a t i u m l i b e r a t i o n , t h e l a t e r a l spermatangial margins begin to erode. This s t a r t s a t t h e apex and p r o c e e d s b a s a l l y i , The remaining a p i c a l p o r t i o n of the t h a l l u s c o n s i s t s of v e g e t a t i v e c e l l s and c a r p o s p o r a n g i a (OBC 5 7 1 8 4 ) .  Figure  5d.,  S p e r m a t a n g i a f o r m i n g m a r g i n a l and s u b m a r g i n a l s t r e a k s which a r e o r i e n t e d i n an apex t o b a s e direction (OBC 5 7 1 8 0 ) .  Figure  5e.  The t i p s o f o l d e r t h a l l i o f t e n have a s l i g h t l y m o t t l e d a p p e a r a n c e c a u s e d by abundant carposporangia (OBC 57192) .  53  Porphyra t h u r e t i i Figure  6a.  M o r p h o l o g i c a l v a r i a t i o n i n t h a l l i w h i c h were g r o w i n g on t h e s t i p e o f N e r e o c y s t i s l u e t k e a n a i n B a r k l e y Sound. Note t h e c h a r a c t e r i s t i c r u f f l e d m a r g i n s (DBC 5 7 2 0 0 ) .  Figure  6b.  The p r o d u c t i o n o f s p e r m a t a n g i a b e g i n s a p i c a l m a r g i n a l r e g i o n o f the t h a l l u s  Figures  Figure  6c  6e.  i n the (UBC 5 2 1 0 9 ) .  and 6d., On o l d e r t h a l l i s h o r t s t r e a k s o f spermatangia form s u b m a r g i n a l l y . Note t h e c h a r a c t e r i s t i c p a t t e r n t h a t r e s u l t s (UBC 57201 UC 95596 r e s p e c t i v e l y ) .  and  Specimen f r o m t h e Monterey P e n i n s u l a w h i c h r e s e m b l e s s p e c i m e n s f r o m B a r k l e y Sound i n s i z e and spermatangial pattern.  54  55  PART I I - SPERMATOGENESIS AND CARPOSPOBOGENES.IS  Introduction  Hus per  (1902)  was  spermatangium  taxcnomic thought the  the f i r s t and  1  criterion  t o u s e t h e number o f s p e r m a t i a  carpospores  for  delimiting  t h a t an u n d i f f e r e n t i a t e d  carpogonium,  and t h a t  Hus as ' c r u c i a t e  1  to  a group  of four  spermatangia.  Hus  devised  division)  spore  and carposporangium  representing  the  A point  has been  the  exception  horizontal  Hus'  notation,  carpospores (Hus, pointed  applied  plate  to  workers,  ( 1 9 7 2 ) , was t h a t  a  to  cube  22, f i g . 25) .  c u t , subseguent  workers  formula  *b'  four  4x4x2,  with  Hus c o n s i d e r e d spermatangia. would  carposporangia  and s p e r m a t a n g i a .  As K r i s h n a m u r t h y  be:  have d r o p p e d  the  I have f o l l o w e d  any i n d i c a t i o n  that  32 c/2  (1972) h a s  1/2  and a p p l i e d t h e f o r m u l a e q u a l l y  could not find  the  *a* and  a/4, b/4, c/2 o r 8 s p e r m a t i a 1/2 a / 2 , 1/2 b / 2 ,  1902,  I  t o be c u b e s w i t h  eguivalent  spermatangium  because  rise  considering  o v e r l o o k e d by a l l r e c e n t  t o be  gave  c o o r d i n a t e s and ' c ' t h e v e r t i c a l .  o f Krishnamurthy  each carposporangium  (referred to  cell  by  He  f u n c t i o n e d as  divisions  formula  as a  species.  vegetative c e l l  c f a vegetative  spermatangium  which  Porphyra  two a n t i c l i n a l  by  a  per carposporangium  this  i n the t o both system  a carpogonium i s  •-Hus (1902) and many s u b s e g u e n t w o r k e r s used t h e t e r n Vpacket' i n p l a c e o f s p e r m a t a n g i u m and c a r p o s p o r a n g i u m . See t e r m i n o l o g y section f o r further discussion.  56  equivalent  t o four  Judging (1956), area  by r e p o r t s i n t h e l i t e r a t u r e ,  of considerable  Jolis  to  According  the carpogonia  to  Drew,  o f P. l e u c o s t i c t a  Berthold  Thuret  l a r g e r and p a l e r t h a n v e g e t a t i v e  carpogonia  Berthold  (1882) f i g u r e d a l s o had s l i g h t  w h i c h d i d n o t push o u t t h e o u t e r  caused  a  considered  these  spermatangium  Drew  Kunieda  not  they  and  to  the  that there  Berthold  He  which  apparently  carpogonia. characterize  Porphyra  formed that  bipolar  by he  species  the he  undifferentiated  could  division  distinguish by  the  was t o o much  wall  variation  be a b l e t o d i s t i n g u i s h v e g e t a t i v e  carpogonia  prototrichogynes Cole  on t h i s  carpogonia Dangeard  f. l i n e a r i s  'trichogyne-like* Based  were  (1956) n o t e d  (1939) r e p o r t e d  umbilicalis  and  of  cells.  from  divisions.  undifferentiated  Conway  wall.  adequately  reported  Porphyra tenera.  with  outer  from s p e r m a t a n g i u m  thickness  reproductive  the  wall.  i n Le  protuberances  t o be o l d u n f e r t i l i z e d  thouqht  division  thickness.  bipolar  carpogonium  cells  vegetative  long  of  did  or  He  vegetative  in  cells  (1902)  studied.  with  protrusion  Hus  wall  cells  Drew  has been an  slightly  also i l l u s t r a t e d  £•  confusion.  by  be  protuberances  in  summarized  r e c o g n i t i o n of the carpogonium i n Porphyra  (1882) c o n s i d e r e d  The  spermatangia.  in  i n P. u m b i l i c a l i s  prototrichogynes  (1927)  P. l i n e a r i s  (Greville)  (1973) d e s c r i b e d  with  Harvey)  illustrated Greville and  (Linnaeus)  'carpogonia-like*  p r o j e c t i o n s i n P. p a p e n f u s s i i  (as  carpogonia J , Agardh. cells  with  Krishnamurthy.  d i s t r o m a t i c s p e c i e s , Conway e t a l . (1975, f i g . 1)  showed t h e c a r p o g o n i u m  with  one p r o t o t r i c h o g y n e .  No  indication  57  was g i v e n o f t h e a p p e a r a n c e species.  Kurogi  monostromatic had b i p o l a r In of  (1961), i n h i s e x c e l l e n t  Porphyra  to the d i f f e r i n g  carpogonium,  conflicting  species,  reports  Drew  two  d i v i s i o n s were a n t i c l i n a l  Krishnamurthy,  1873; 1927;  periclinal  Kurogi,  could  (Janczewski, Hus,  1902;  Krishnamurthy the  little repeated  considerable  1873;  Thuret  Ishikawa,  whereas  1924;  did  Dangeard,  that the  not  initial  d e p e n d i n g on indicate  amount  at  the  and  1921;  of  the  light  observation microscope  Grubb,  1924;  and  than  in  division  been  well  indicate of that  the  sequence  that  Tanaka,  1952; 1975),  These  studies  of did  t h e s p e r m a t i a a r e formed by  spermatanqium.  Janczewski  1882;  o f the spermatangium  documented.  of  level  B o r n e t , 1878; B e r t h o l d ,  1959, 1972; Humford, 1973a; Conway e t a l .  division  contradictory  1902;  (Janczeuski,  or p e r i c l i n a l  a l . (197 5)  other  o f the carposporangium.  has n o t more  et  first  of t h e carposporangium  (1952) i n d i c a t e d  anticlinal  development  appearance  Por£hyra  Tanaka  the  1964b),  I s h i k a w a , 1921; Grubb,  be e i t h e r  a  spermatangial  division of  (Hus,  ( i n the plane of the t h a l l u s )  Conway  Despite  Conway,  division  there are  a n g l e s t o each  divisions)  1972;  the f i r s t  1882;  sequence  cruciate  1969b,  1961).  species.  division  4  appearance  pointed out that  and a t r i g h t  as  1959,  have r e p o r t e d  division the  to  Berthold,  r e p o r t s about the  One g r o u p o f w o r k e r s b e l i e v e d  referred  as b e i n g  of  t h a t t h e carpogonium  of the o r i e n t a t i o n o f the f i r s t  carposporangium.  others  investigation  reported  (1S56)  the  (often  i n monostromatic  prototrichogynes.  addition  the  o f t h e carpogonium  (1873),  The  data  are  T h u r e t and B o r n e t  58  ( 1 8 7 8 ) , Hus  (1902), Ishikawa  that the f i r s t whereas  division  Tanaka  anticlinal. discrepancies  It  was  statement,  Drew are  distinguishing  the  to  me  normal  has  the  reported  that  was  genus  valid.  preliminary  that  of  manner.  formation therefore  in  These  described.  595)  and  to t r y to  follow  microscope  of  spore  decided  and  their  level  Hollenberg  by  a  monospore  for  at  the study s i t e  confirm  that  curving  were r e l e a s e d  and  t h e genus  and  in  site  forming  description  at t h a t t i m e )  cell  singly.  the study  were n o t  f i t the  gardneri  (1943)  division  A d e t a i l e d s t u d y o f carpogonium  essential to  incorrectly  from  carpospores  specimens  ( Porphyrella  plants  p.  t o t h e method  specimens from  in  respects.  formed of  t h a t t h e c a r p o g o n i a and  ether  in  cell.  (1956,  been p a i d  that the carpospores  examination  £oj£hyra g a r d n e r i  various  difficulty  Drew's  Smith  was  indicated this  the  i t was  studying.  to  and  periclinal,  a vegetative  light  maintained  these  carpogonium  at the  the carpogonium  Erythrotrichia.  out,  I therefore  Porphyrella  similar  was  cell-division  still  species I  (1924)  (1959) t h o u g h t  to  that  t h e s p e r m a t a n g i u m and  For  My  perhaps  seguence i n d e t a i l  division  pointed  attention  was  characterize  the t h r e e  due  "Little  formation..."  Krishnamurthy  (1956)  apparent  Grubb  spermatangium  spermatangium from  distinguishing  division  of the  (1952) and  as  (1921) and  in  of a l l  carpospore  type l o c a l i t y  was  Porphyrella  was  59  Materials Spermatangial the  study  (approx,  glutaraldehyde  hours.  and  were  2  filtered  and c a r p o s p o r a n g i a 1 t h a l l i  site  5mm )  Whenever  marginal  cut  (50%),  seawater  And Methods  out  and  phosphate  (1:4:4) possible  and  submarginal  fixed buffer  (McBride  and  m a t e r i a l was  were c o l l e c t e d a t  in  a  soluticn  pH 7.2, and Cole,  fixed  pieces of  millipore  1969)  for  immediately  1.5  i n the  field. Fixation series  was  f o l l o w e d by d e h y d r a t i o n i n a g r a d e d  and i n f i l t r a t i o n  JB-4).  Serial  sections  glass knives using a stained water  with  with g l y c o l (Henry,  Sorval  tcluidine  ( F e d e r and O ' B r i e n ,  1977) 2-3  JB-4  microscope  Photographs automatic  (Model  were t a k e n  photographic  um  (Polysciences  thick  microtome.  i n benzoate  buffer  1968)  mounted i n  Euparal  a  attachment  or  Nikon  Microflex  were  distilled (GBI  and examined on an  FHA) w i t h p l a n a p o c h r o m a t i c with  were c u t c n  Sections  blue-0  L t d . , Heaton S t , Denton, Manchester) light  methacrylate  ethanol  Labs  Olympus  objectives. (Model  AFM)  u s i n g Kodak Panatomic-X  film.  60  Besults  A.  Porphyra  1.  gardneri  Spermatogenesis  The  division  spermatangia section. central  was  Vegetative  During l a t e  of  to  and  each  was  third  mature was  formed  surface Figure  view 7i.  2.  periclinal  7d  periclinal  The  (a/1,  in  This  and  containing  Figure division  and  7a).  along  the  cells  went  first  b/1,  8a).  64  the  a  production  division  c/2)  of  the  (Figure 7c).  The  and  These  at r i g h t  (Figures  7e-f)  spermatia  (a/4,  mature  angles  were f o l l o w e d  immature s p e r m a t a n g i u m  7h,  (Figure  that vegetative  divisions  An  transverse  forming  t o a s p e r m a t a n g i u m was  (Figures  mature  by  until b/4,  c/4)  i s shown  in  spermatangium  s e q u e n c e i s summarized  in Figure  a  in  8a.  Carposporogenesis  The  spermatangia,  ( F i g u r e 9b)  are  surface  (Figure  By  nucleus  began  stage  ( F i g u r e 7b) ,  (Figure 7g).  and  of  have a s i n g l e s t e l l a t e c h l o r o p l a s t ,  first  spermatangium  formation  view  d i v i s i o n s were a n t i c l i n a l  other  a n t i c l i n a l and  surface  spermatangia  The  wall layer  in  to the  peripherally located  transition  spermatangium second  and  margin.  a new  cells  A p r i l the  t h r o u g h i n the  leading  followed  pyrenoid,  thallus  sequence  view early  May  carpogonia  and  r e a d i l y d i s t i n g u i s h e d from  carposporangia  vegetative  cells  in  9a).  carpogonia  began f o r m i n g i n  the  submarginal  61  region  of  the thallus.  No c u r v i n g , E r y t h r o t r i c h i a - l i k e  were i n v o l v e d i n t h e i r f o r m a t i o n . carpogonia their  were  characteristic  presence  of  carpogonium and  carpogonia  formed of  were c l o s e l y from  starch  lemon-shape  carpospores that  from  cells.  per  carposporangium  the  carpospores  a single  In  to  before a  This  appeared  immediately  thick  was  The  c a s e s two  wall  In determining i t  the  pyrenoid,  some  differentiation  division  the  (Figure 9c).  (Figure 9d).  carpogonium  t h e daughter  due  chloroplast,  grains.  appressed  a vegetative c e l l  ascertain  or  section  v e g e t a t i v e c e l l s by  prototrichogynes  floridean  between  derived  spindle-  from  contained a single s t e l l a t e  result  following  transverse  distinguished  bipolar  abundant  to  easily  In  nails  had  t h e number  necessary  being observed  were  c a r p o g o n i u m and n o t two c l o s e l y  to  actually appressed  ones. Following without first c/2)  i t  fertilization  in  division  some  this  9h). occur  most  carpogonium  stage,  In  was p e r i c l i n a l  view  were  form  4  underwent  did  (a/1, b/1, not  second  carpospores  divide  and  third  ( F i g u r e s 9 f , 9g,  two d i v i s i o n s  ( a / 1 , b/2, c / 2 . F i g u r e 8 b ) ,  carposporangia  ( F i g u r e s 9g 8  perhaps  appeared but  o r i e n t e d a t r i g h t a n g l e s t o each  The l a t t e r  t o produce  (Figures 9 i  some  some c a r p o s p o r a n g i a t h e s e  they  (Figure 8b).  division  to  i n t h e same p l a n e  cases  surface  tut  (or  t o be d i s c u s s e d i n P a r t I V ) , t h e  of the carposporangium  anticlinal divisions  to  cases;  the  ( F i g u r e 9 e ) . , Many o f t h e c a r p o s p o r a n g i a  beyond  and  of  and 9h)..  carpospores  and 9 j ) ,  The  were  easily  Occasionally  (a/2, b/2, division  c/2)  in  other  seen  in  further occurred  sequence  of  62  c a r p c s p o r o g e n e s i s i s summarized  3.  Examination  i n F i g u r e 8b.  Of Type L o c a l i t y  Plants  In view o f t h e mode o f c a r p o g o n i u m f o r m a t i o n a n d sequence it  of  the carposporangium  was a p p a r e n t  t o me t h a t t h e y  not  Porphyrella.  of  herbarium  revealed  specimens  that  Therefore  they  locality  the  taxon.  same From  the notes  herbarium,  belonged  were  i n study  Alaska  and  to  identical  to  t o determine  made  i t was  description,  by  study  characteristics  marginal  distribution  i n common:  on  spermatia/spermatangium, s i m i l a r  basal  plants.  California,  on  a  in his  t a x a had t h e  host  monospore  region  were  s p e c i m e n s , and  specimens  similar  Laminaria.  California  site  p o s s i b l e t o c o n f i r m t h a t both  following  Porphyra,  whether t h e p l a n t s a t  herbarium  Dr. H o l l e n b e r g  plants  carposporangia  northern  on t h e M o n t e r e y P e n i n s u l a ,  original  site  t o t h e genus  of carpogonia  from-  i t was n e c e s s a r y  the type  thorough  Examination  observed  division  plants,  a  cycle,  64  morphology  and  a t t a c h m e n t , v e g e t a t i v e felade t h i c k n e s s 25-35 a n , and v e g e t a t i v e thalli  minutely  locality reported  wrinkled.  p l a n t s and t h o s e mode  of  The  from  o n l y d i f f e r e n c e between  the rest  carpogonium  of the  formation  and  coast  was  type the  carposporangium  division. In order in  July  Mission  to solve this  problem, specimens  were  1976 a t P o i n t J o e ( t y p e l o c a l i t y ) ,  Pescadero  Point,  of  carposporangia however, t h e i r  California. was  The  number  and  division  P o i n t and  carpogonia  l o w compared t o more n o r t h e r n  mode o f f o r m a t i o n  collected  and  populations;  was  identical  63  with  that  observed  i n specimens from  areas along the coast. culture  and  the  same  T h i s o b s e r v a t i o n combined  chromosome  type l o c a l i t y  as  and o t h e r  with  field,  s t u d i e s ( s e e P a r t s I , I I I , I V ) on t h e  population conclusively  taxon  the study s i t e  was  found  demonstrated  at  the study  that  site.  i t  was  I t seems  p r o b a b l e t h a t t h e s m a l l number o f c a r p o g o n i a a n d c a r p o s p o r a n g i a produced their  B.  by t h a l l i a t t h e t y p e  the  reason  that  follow  in  Porphyra n e r e o c y s t i s  Spermatogenesis  Spermatogenesis Porphyra  wall  from  was l a i d  division. series  vegetative down a r o u n d  Furthermore,  difficult  to  spermatangia  cells.  were  not  U n l i k e P. j a r d n e r i ,  t h e spermatangium the spermatangia  prior  In  t h e number o f s p e r m a t i a  s u r f a c e view.  distinguish several  The  from  divisions  Spermatangia spermatia  vegetative  (Figures  b/4,  difficult  spermatangia.  o f t h e t h a l l u s i s shown  spermatangia cells  first  were  until  they  difficult  to  had u n d e r g o n e  (Figure 91). with  4 ( a / 1 { 2 ) , b / 2 ( 1 ) , c/2) o r 8 ( a / 2 ,  were s t a r t i n g t o l o s e p i g m e n t a t i o n .  divisions occurred (a/4,  young  no new  t h a t h a d undergone a  i n mature  F i g u r e 9k a v e g e t a t i v e p o r t i o n  very  to i t s  o f d i v i s i o n s were n o t w e l l d e f i n e d , making i t  determine  c/2)  was  n e r e o c y s t i s because the  distinct  in  is  f o r m a t i o n has b e e n o v e r l o o k e d i n t h e p a s t .  1.  to  locality  c/8)  ( F i g u r e 10a) u n t i l containing  10b and 1 0 c ) .  128  a  mature spermatia  T h i s i s a maximum number;  b/2,  Subseguent  spermatangium was  reached  variations in  64  the  final  2.  number o f s p e r t a a t i a / s p e r m a t a n g i u m  the  form  characterized section  region  pyrenoid  the t h a l l u s  difficult  thallus.  The  In t r a n s v e r s e because  carpogonia  i t were  transverse  compared t o t h e s q u a r e - s h a p e d  vegetative  They had a  single  stellate  recognized  fertilization  Part  January  in  c o u l d a l s o be  in  in  to recognize  and p e r i p h e r a l n u c l e u s .  forming  discussed  the  as b e i n g r e c t a n g u l a r and e l o n g a t e  ( F i g u r e 10d).  spermatia  was  of  prototrichogynes.  ( F i g u r e 10e)  central of  c a r p o g o n i u m f o r m a t i o n began  t h e carpogonium  not  cells  site  submarginal  section did  observed.  Carposporogenesis  at t h e s t u d y in  were  IV)  and  chloroplast,  Carpogonial by  the  regions  presence  canals to the carpogonia by  periclinal  divisions  of  ( t o be of the  carposporangia. Frequently (Figure cf  10f),  t h e number The  (a/1,  nornally and  of carpospores division  c/2)  b/2,  c/2)  at  this  the »c -axis ,  that  periclinal  this  Both  and a t r i g h t  division  the  second  a n g l e s t c each  at this stage  (Figure 10i).  stage  was  (see P a r t I V ) .  were  Subsequent  carposporangia  (see P a r t I I I ) .  of t h e carposporangium  Further  easily  division  8 carpospores/carposporangium  ( F i g u r e s 10h and 1 0 j ) . Host  carpospores alcng  It is felt  Carposporangia  t o produce  observed  to misinterpretation  o f t h e carposporangium  d e t e c t a b l e i n s u r f a c e view rapidly  lead  were  carposporangium.  were a n t i c l i n a l  ( F i g u r e 10b).  occurs  per  ( F i g u r e 10g) .  divisions  carpogonia  w h i c h may  follows f e r t i l i z a t i o n  third  other  appressed  a phenomenon  first  b/1,  2 closely  (a/2,  released division  ( F i g u r e 101) o c c u r r e d  65  frequently  on  older  pigmentation. divided mixed  the  was  unable  fa/4,  Porfhysa  have  2 closely  to  was  not  detail  as  sequence  incompletely  pigmentation  and  were  gave t h e t h a l l u s  b/2  and  a  a/4,  that  b/2  a  reported  1944;  and  11a)  single  this. b/4  Conway e t appeared  were t o o  large  carposporangium.  although  I  for  Krishnamurthy,  superficially  t o form  a/4,  carpospores  H o l l e n b e r g , 1976;  from  r e p o r t e d t o be  32  typically  ( F i g u r e s 10k  appeared  with  Smith,  I  suspect  are actually  could  not  that  the  made up  of  carposporangia.  thuretii  a limited  possible was  amount o f m a t e r i a l o f P o r p h y r a  to  done  follow for  spermatogenesis  P. g a r d n e r i .  i s shown i n F i g u r e s 11b  and  defined i n transverse section,  determine prior  deeper  Spermatogenesis  Due  well  Abbott  determine  appressed  Porphyra  1.  been  1902;  derived  a/4,  carposporangia  it  has  formula  been  uneguivocally  a  and  their  carposporangia  as  1973a;  spore  Carposporangia  cells  lost  G r o u p s of c a r p o s p o r e s  t o have t h i s  C.  to f i n d  n e r e o c y s t i s (Has.  1975).  to  vegetative  in  (Figure 5e).  c/2)  Mumford,  resulted  mature c a r p o s p o r a n g i a  appearance  (a/4,  1972;  and  c a r p o s p o r a n g i a w h i c h had  mottled  al.  Suspected  i n with  I  thalli  tc  i f this  i s due  to the  11c. but  The The  in  thuretii the  early  same  division  spermatangium i s  I have not  formation of a  new  i n t e r m e d i a t e stage  in  been a b l e t o wall  layer  division.  Figure  11d  shows  an  spermatangial  66  divison. c/4)  Mature s p e r m a t a n g i a c o n t a i n e d  (Figures  2.  not i*«  11e and 1 1 f ) .  transverse  seen  This  to  section the carpogonia  possess  fixed  until  9§£<J g£i  Porphyra  prototrichogynes  f o r good c a r p o g o n i u m  degree  therefore The  2 days a f t e r being  of  thuretii  ( F i g u r e 11g).  than  stages). (a/2, release  division observed  in  (Figures  P. g a r d n e r i  The mature c a r p o s p o r a n g i u m  b/2,  c/2)  could  occur  carposporangium. a mottled,  mature  a single  ( a / 1 , b/1,  sequence  (Figures  11k  deeply  carposporangia.  distal pigmented  i n the f i e l d  was  preservation.  stellate  may  c/2)  division of  f e r t i l i z a t i o n ; see (Figure  11h).  The  1 1 i and 1 1 j ) was t h e same and  P. n e r e o c y s t i s  contained  and 1 1 1 ) ,  region  chloroplast,  The f i r s t  8  of older  fearlv  carpospores  although  any t i m e a f t e r t h e f i r s t The  of  reported.  (following a suspected  P a r t I V ) was p e r i c l i n a l  was  studies  d e v e l o p m e n t i n P. t h u r e t i i  and p e r i p h e r a l nucleus.  carposporangium  that  I have  carpogonium c o n t a i n e d  subseguent  My  and s p e r m a t a n g i u m  prototrichogyne  be g r e a t e r  c e n t r a l pyrenoid the  collected.  i n d i c a t e d t h a t immediate f i x a t i o n  n  The  had  slight  of  needs t o be r e - e x a m i n e d b e c a u s e t h e m a t e r i a l I s t u d i e d  essential  as  ( a / 4 , b/4,  Carposporogenesis  In were  64 s p e r m a t i a  carpospore  division thalli  a p p e a r a n c e due t o t h e  of  the  typically abundant  67  Discussion  This yielded  study  three  of  major  s p e r m a t o g e n e s i s and  The  genus P o r p h y r e l l a i s n o t  2.  An  attempt  has  carpogonium The  and  division are not released  by  of  the  number o f  division  pattern  the  genus  Hollenberg nomenclatural  The  genus  g a r d n e r i . Kurogi  this,  the  i n the  mode  as  Porphyra  of  seguence  of  the  proposed  by  Hus  s p e c i e s examined.  carpogonium  formation  (194 3)  carposporangial (1961) has for  Porphyrella not  c h a n g e was  Porphyra g a r d n e r i Synonym:  made  (Smith  kuniedai as  valid.  Description:  chloroplast  per  and  unique the  same  I n view  by  Smith the  to  of and  following  (Hawkes, 1977b):  et  Hollenberg)  J . B o t . , 30:  Thalli  not  Kurogi.  Therefore  not  small  figured  described  Porphyrella gardneri  ( 1 9 4 3 ) , Am.  and  are  seguence i s  the  d i v i s i o n s are  reported  Porphyra  is  division  Furthermore,  and  carpogonia  the c a r p o s p o r e s  carposporangial  Porphyra.  the  Porphyra.  spermatangia  occur  characterize  division  u n e g u a l d i v i s i o n s and  singly.  variable S»  to  i n Porphyra g a r d n e r i demonstrated t h a t the formed  typical  and  does n o t  of  made  *cruciate*  carposporangia  Examination  been  valid.  spermatangium of  classical  (1902)  has  results:  1.  3.  carpcsporogenesis  comb.  Smith  nov.  et  Hollenberg  215-216, f i g s .  13-14.  m o n o s t r o m a t i c 25-35 um  thick,  cell.  Thalli  up  to  130  mm  one long  68  and  60 mm  10-30  wide,  mm  wide.  monospores. spermatia formed and  but t y p i c a l l y Reproducing  Mature  ( a / 4 , b/4,  by  undergoing  d i v i s i o n s t o form  (a/1(2),  released  The  phase.  have b e e n e x a m i n e d 1970; C o l e ,  formation  the  basis  of  Smith  occurrence  of  both  Porphyra  aardneri.  affinities  with  spore  division spore  phase  a  with p i t  species  that  1970; B o u r n e e t  formation  of t h e  flollenberg's modes  by  repeated  Bangiaceae.  (1943)  of  spore  (1956) s u g g e s t e d  of the  i s c h a r a c t e r i s t i c of  spermatia/spermatangium  i s typical  families.  P. g a r d n e r i i s t y p i c a l  into  197 2b) .  and  Drew  both  germinating  several  these  c/2)  i s heteromorphic,  (Lee and F u l t z ,  t h e E r y t h r o p e l t i d a c e a e , whereas  carpospores/carpospcrangium  ( a / 2 , b/2,  o f a l l Porphyra  by u n e q u a l  produce  cell  prototrichogynes.  Conchocelis  plugs as i s t y p i c a l  to  history  carpospores  conchocelis  division  Carpogonium  ( a / 1 , b / 1 , c/2) , f o u r  eight  life  64  one t o t h r e e o r r a r e l y  two  b / 2 { 1 ) , c/2) o r  carpospores.  Spore  maximum.  bipolar  by  containing  by t h e t r a n s f o r m a t i o n o f a v e g e t a t i v e  Carposporangium  al.  c/4)  l o n g and  asexually  spermatangia  characterized  seven  20-70 mm  cr On  r e p o r t of the formation  that t h i s  in  s p e c i e s had  T h i s , however, i s n o t t h e c a s e ; Bangiaceae  in  a l l aspects  of  formation. Only  one  other  £2I£hyxe.11 a, t h i s described  by  species  being  Hcllenberg  has  been r e f e r r e d  t o t h e genus  californica  Hcllenberg,  Porphyrella (1945)  from  Santa  C r u z I s l a n d and  69  several He  places along  separated  saxicolous  i t  thallus of  from  holotype  i n carpogonium  are  probably  carpogonia seen.  P o r p hy r e 1 l a  ca1ifornic a  until  c o n c l u s i v e c a n be s a i d are  view  should  further  An  examination  the thallus  No e v i d e n c e  of this be  margin. which  o f uneven  i t i s suspected  transferred  investigation  about  at  species.  were l a r g e r p i g m e n t e d c e l l s ,  In  of  (GJH 1353) c o n s i s t s o f one  and c a r p o s p o r e s .  was  There  i n this  on a g l a s s s l i d e .  region  crosswalls  however,  No m e n t i o n was made  abundant spermatangia a l o n g  Also i n the marginal  California.  on t h e b a s i s o f i t s  formation  o f P. c a l i f o r n i c a  p e r m a n e n t l y mounted  i t revealed  southern  P o r p h y r a garjinjarj.  h a b i t and shape o f t h a l l u s .  uneven c r o s s w a l l s The  t h e mainland c o a s t ,  to  that  Porphyra;  o f i t i s made,  nothing  i t s taxonomic s t a t u s .  least  three  different  carpogonial  m o r p h o l o g i e s i n t h e genus P o r p h y r a .  Hawkes (1977b) p o i n t e d o u t  that  prototrichogynes  in  those  reported, species  species  there  are  for  two  have c a r p o g o n i a  in  distromatic  prototrichogyne, by  produced  specialized  types.  species  the  carpogonium  have  l o c a t e d on t h e  surface, has  whereas only  l o c a t e d next t o the t h a l l u s s u r f a c e .  with by  (Table  V I ) , most  prototrichogynes,  the  receptive  fertilization  (see  characteristic  elongate  developing sites  Part  Judging species  prototrichogynes  carpogonia  and f u n c t i o n as  They  give  lemon-shape and cause  wall of the t h a l l u s t o protrude  Porphyra  one  The  f o r spermatium  IV),  been  Moncstrcoiatic  2 prototrichogynes  next t o t h e t h a l l u s  r e p o r t s i n the l i t e r a t u r e  have c a r p o g o n i a are  carpogonial  with  s i d e s o f t h e carpogonium  which  to varying  attachment the the  and  carpogonia cuter  a  cell  d e g r e e s d e p e n d i n g on  70  how  well developed  to  vary  £•  tJLSsi  Table  d e p e n d i n g on  VI  of  long  The  J o f f e ' (1896) t o  third  The  type  shape.  of  carpogonial  to  specifically  prototrichogynes s p e c i e s from the lack  E.  M.  Berthold  level i s  (1882)  carpogonia  prototrichogynes  slightly  Dangeard noted  in  and were  morphology i s d i s p l a y e d  differentiate  cells i s their  Only  listed  gardneri.  feature  vegetative  at the by  and  prototrichogynes.  old unfertilized  n e r e o c y s t i s i n which no  only  them  by  reported  papenfussii species  long  prototrichogynes  formed  seen i n P o r p h y r a  Porphyra  extremely  protuberances  be  degree of p r o t r u s i o n appears  u n i q u e amongst t h e  their  these  The  species.  because  needed.  Porchyra  the  Ueda a r e  of  ft  are.  (Yendo)  Examination  not  they  more  (1927) and  that  a  Northeast  formed.  carpogonia  elongate,  T s e n g and  few  (Table VI).  the  are  (1955) have  species  Preliminary observation  P a c i f i c Ocean s u g g e s t s  prototrichogynes.  Further  from  rectangular  Chang  Porphyra  by  lack  of  other  t h a t some  investigation  of  of t h i s  is  needed. Dangeard  (1927) was  species  possessed  (Table  VI).  the  Tseng  and  Chang  i n the  Porphyra  species  Porphyra  s p e c i e s have t h e  that t h i s  may  that  cell those  ability  to  t o r e p o r t t h a t some P o r p h y r a  prototrichogynes,  variation  tough  first  occurrence they  species  the  (1955)  ability  noted  They  which  Another e x p l a n a t i o n  them  three  that  all  prototrichogynes possess  which I am  was  i n the  suggested  t o form  i n those  lacked  that there  prototrichogyne  studied.  be s u p p r e s s e d wall.  of  whereas o t h e r s  a  but  thick,  proposing  is  lacking  prototrichogynes  have n e v e r had  the  produce these  more s p e c i a l i z e d  receptive areas  and  71  may t h e r e f o r e other  by more p r i m i t i v e .  Porphyra s p e c i e s  carpogonium o f t h i s Groups  of  P. g a r d n e r i resulting  from  reported  Conway  appears  The  laid  from  down  although  making  to  In  nereocystis  to  that  gairdneri  cells  not  cell  seen.  as This  P. p a p e n f u s s i i  by t h e new  and  adding  the  more  more s p e c i a l i z e d  study suggests that  the  note with  that the  that  spermatangia, i t f o rthe The new  appearance,  p r o d u c e d by one  spermatangium  wall  layer  single this  evidence  making  i t  specialized  carpogonium  in  support  c f the In  advanced carpogonium i s a s s o c i a t e d  with  may  Thus,  the  of  genus.  spermatangium.  of  of  spermatangium.  less  simple  be  i s the f i r s t  defined  the new  layer  reported  a l l the spermatia  further  there  and  This  can  (Harvey) K y l i n .  the products of a to  wall  f o r Porphyra  i t i s a more p r i m i t i v e member  carpogonium  in  of 4 carpogonia  Porphyra g a r d n e r i  contrast,  i s associated  P. n e r e o c y s t i s .  of  were  hypnoides  lacked  interesting  spermatangium  noted  vegetative  (1973a) p r e v i o u s l y  to identify  It  the  in  Ptilota  to identify  £•  limited  has been r e p o r t e d  difficult  theory  of a  a l . 0975)  be  groups  g i v e s t h e spermatangium a w e l l  spermatangium. Porphyra  in  c h a r a c t e r i z e the  occasionally  but  division  vegetative  of  i t simple  i s  et  and D i x o n  tetrasporangia layer  were  around i t p r i o r t o d i v i s i o n .  Scott  studied  Hollenberg.  time t h i s f e a t u r e  wall  1  spermatangium  distinguished  t o completely  P. n e r e o c y s t i s .  P. s c h i z o p h y l l a  is  carpogonia  a •cruciate  by  phenomemon  i n order  needs t o be  genus.  2  and  This  this  preliminary  be a c o r r e l a t i o n between t h e t y p e  spermatangium. / Study o f o t h e r  members o f  72  t h e genus i s n e e d e d t o c o n f i r m The  report  carpospores This  may  not o b s e r v e d  the  recognized, 3  the  occurred. different  Hus (1975)  Hus It  (1902) is  to  larger  the  surface  contact  resembles detected  spermatangia  division  those  and  The  first  as  first  Krishnamurthy  the  in  (1972),  (1959)  seguence  is  should  be  c a s e s where  and  cruciate  of b/2,  this  c/2).  size.  c/2-4, but first  Conway  of Porphyra  (a/4, b/4,  Porphyra  a/2,  b/2  carposporangia  this  be f o l l o w e d .  were i d e n t i c a l .  by  been  On  I  et  al.  nereocystis was  unable  older t h a l l i  they  ;  were  I  never  r e p o r t of carpcsporangia  i n Porphyra n e r e o c y s t i s .  ( K u r o g i , 1972).  a  have  s p e c i e s : however, t h i s  T h i s i s the  Japanese  of 2  Such a  •cruciate* divisions  that  carpospores  dimensions b/4  Janczewski.  the  mature c a r p o s p o r a n g i a  this.  t h a n c/2  enigma.  reported.  carposporangia  than  a/4,  later  c a r p o s p o r a n g i a a/4,  In  of  and  Krishnamurthy  up t o 32  identify  larger  no  particularily  reported  observed  and  possible  have been  by  and  an  seguence can  both  divisions  f o r other Porphyra  (1902),  contained  spermatia  spermatangium  division  for  the i n i t i a l  re-investigated, divisions  and  periclinal  by  and  subseguent  species  is  reported  of  i n the 3 s p e c i e s I s t u d i e d .  carpogonium  carpcsporangia division  (1873)  h a v e been a m i s i n t e r p r e t a t i o n  Once  a l l  Janczewski  o c c u r r i n g i n t h e same s p o r a n g i u m r e m a i n s  phenomenon was  In  by  this.  single  species  o r a/4, Kurogi  b/2;  carposporangia there are  (1972) n o t e d  often  carposporangium  phenomenon by c o m p a r i n g  occurs of  and a/4,  no  have  reports  that the  close  superficially b/4.  the dimensions  Kurogi of  these  73  large  'carposporangia*  vegetative  cells.  with those of s m a l l e r c a r p o s p o r a n g i a  They  however, t h e s u s p e c t e d to  be d e r i v e d from I  agree  should  a/4,  b/4  a single  •carposporangia*  (1972)  responsible  f o r the r e p o r t s of carposporangia the N o r t h e a s t  have c a r p o s p o r a n g i a MaBlJSggagis 1902;  Conway  as a/4,  b/4  et  (Mumford,  or  (1961)  the  that  could that  division  numbers  transverse this  cause  of  section  accepted  spermatangium certainty.  unless or  Other  Porphyra  perforata  <Hus,  should as  be large  of  such  some v a r i a t i o n  in the  sporangium.  Kurogi  per  the  thallus  variation.  spermatangia  put  on  as  those  and  with (1972)  the d i v i s i o n taxonomic  t o emphasize the  products  carposporangium  and  Furthermore, g e n e r a l l y had  thin  thalli.„  suggested formulae  criteria, et  of  he larger  view o f  that  less  spermatangia  a l . 1975).  of the d i v i s i o n be  In  in  a t r e n d which i s  that the d i v i s i o n  can  ecological  carposporangia  (Mumford, 1973a; Conway  a d d i t i o n I would l i k e meaningless  been  i f carposporangia  with t h i c k t h a l l i  than  carposporangia  generally  age  Krishnamurthy  e m p h a s i s s h o u l d be and  has  b/4.  Carposporogenesis  carpospores  species  variation,  the  occur.  number o f s p e r m a t i a  observed  a/4,  and P.  I n a l l 3 s p e c i e s I s t u d i e d t h e r e was  conditions  large  that  f o r example,  1973b)  i n t h e s e s p e c i e s t o see  noted  think  carposporangia  dimensions,  a l . 1975).  really  were t o o  P a c i f i c Ocean have been r e p o r t e d t o  of these  Mumford  re-examined  appressed  and  of  from  closely  view  phenomenon  species  dimensions;  carpogonium.  with Kurogi*s 2  a l l have s i m i l a r  and  In  formula i s of a  single  recognized  with  T a b l e V I . Summary o f t h e s p e c i e s c f P o r p h y r a w h i c h p r o t o t r i c h o g y n e s , and t h o s e w h i c h do n o t . Taxon Jlthout  possess  Reference  prototrichogynes  1. M o n o s t r o m a t i c Porphyra dentata Kjellman* 2* l i n e a r i s G r e v i l l e E*  With 1.  n e r e o c y s t i s Anderson  T s e n g and Chang (1955) Dangeard (1927) - a s P. u m b i l i c a l i s f» l i n e a r i s ( G r e v i l l e ) Harvey Hawkes ( t h i s t h e s i s )  prgtotrichojyjges  Honostromatic P o r p h y r a a n g u s t a Ueda P. c a p e n s i s K i i t z i n g emend. Agardh E* c r i s p a t a K j e l l man P. g a r d n e r i (Smith e t H c l l e n b e r g ) Hawkes P. k a t a d a i M i u r a P. k u n i e d a i K u r o g i P. l e u c o s t i c t a T b u r e t i n Le J o l i s £• p e r f o r a t a  J . Agardh  £• S i l i o i M S i i i a t a K j e l l m a n z P. t a n e q a s h i m e n s j s S h i n m o r a £. t e n e r a K j e l l m a n P. t e n u i p e d a l i s M i u r a £. u m b i l i c a l i s (Linnaeus) J . Agardh P. y e z o e n s i s Ueda P. s p . ( a m e t h y s t e a Kiitzing?)  Kurogi Graves  (1961) (1969)  T a n a k a (1952) Hawkes (1977b, and thesis)  this  M i u r a (1968) K u r o g i (1961) J a n c z e w s k i (1873) B e r t h o l d (1882) B o s e n v i n g e (1909) Kornmann (1961a) Dr. T. F. Mumford ( p e r s o n a l communication) Tanaka (1952) Shinmura (1974) I s h i k a w a (1921) K u n i e d a (1939) K u r o g i (1961) M i u r a (1961) Dangeard (1927) K u r o g i (1961) Yabu and T c k i d a (1963) Kornmann (1961b)  Distrcmatic i 2 £ 2 h l £ § b u l b o p e s (Y en do) Okamura P. fiapenfjgssii Krishnamurthy  Tanaka  (1952)  K r i s h n a m u r t h y (1972) Conway and C o l e (1973) Conway e t a l . (1975)  75  P. s c h i z o p h y l l a i n Smith and  Hollenberg Hollenberg  P. t a s a  Ueda  (Yendo)  K r i s h n a m u r t h y (1972) as J . n o r r i s i i , Krishnamurthy Conway e t a l . (1975) N a g a i (1941) Tanaka (1952)  »Tanaka (1952) f i g u r e s p r o t o t r i c h o g y n e s f o r t h i s d o e s n o t m e n t i o n them i n t h e d e s c r i p t i o n .  s p e c i e s but  T s e n g and Chang (1955) r e p o r t e d t h e y c o u l d n o t f i n d prototrichogynes i n t h i s species. 2  76  PojEkXXI g a r d n e r i - s p e r m a t o g e n e s i s , l i g h t microscopy Figure  7a.  V e g e t a t i v e c e l l w i t h l a r g e c e n t r a l p y r e n o i d an s i n g l e s t e l l a t e c h l o r o p l a s t . The n u c l e u s i s peripherally located.  Figure  7b.  New w a l l l a y e r (arrow) l a i d down a r o u n d spermatangium p r i o r t o the f i r s t d i v i s i o n .  Figure  7c.  The f i r s t d i v i s i o n periclinal.  Figures  7d-f.  o f t h e spermatangium i s  Subseguent d i v i s i o n s  of t h e  Figure  7g.  Mature spermatangium transverse section.  Figure  7h,  S u r f a c e view o f an immature  Figure  7i.  Surface  spermatangium.  4 spermatia  deep i n  spermatangium.  view o f a mature s p e r m a t a n g i u a .  77  78  Porphyra  gardneri  F i g u r e 8a.  S p e r m a t o g e n e s i s . The d i v i s i o n s e g u e n c e l e a d i n g t o a mature s p e r m a t a n g i u m c f 64 s p e r m a t i a . Note t h e p r o d u c t i o n o f a new w a l l l a y e r ( i n d i c a t e d by a t h i c k e r l i n e ) p r i o r t o d i v i s i o n , and t h e a b s e n c e o f any * c r u c i a t e ' d i v i s i o n s .  Figure  C a r p c s p o r o g e n e s i s . D i v i s i o n seguence which p r o d u c e d 2, 4 o r 8 c a r p o s p o r e s p e r carposporangium. The u p p e r c a r p o s p o r a n g i u m c o n t a i n i n g 4 c a r p o s p o r e s i s most o b v i o u s i n s u r f a c e view and i s shown i n F i g u r e s 9g and 9h. The d i v i s i o n pathway i n d i c a t e d by s o l i d a r r o w s i s most common.  8b.  79  4  zCTI  b  80  Porphyra  gardneri -  carposporogenesis  Figure  9a.  Surface  view o f v e g e t a t i v e  cells.  Figure  9b.  S u r f a c e v i e w o f s p e r m a t a n g i a (lower c a r p o g o n i a and carposporangia.  Figure  9c.  T r a n s v e r s e s e c t i o n o f t b e t h a l l u s s h o w i n g two carpogonia. Note the b i p o l a r p r o t o t r i c h o g y n e s a t t a c h e d spermatium (arrow).  left),  and  Figure  9d.  T r a n s v e r s e s e c t i o n s h o w i n g two c l o s e l y a p p r e s s e d carpogonia. Note a t t a c h e d s p e r m a t i u m (arrow).  Figure  9e.  T r a n s v e r s e s e c t i o n showing t h a t the f i r s t of the caposporangium i s p e r i c l i n a l .  Figure  9f.  T r a n s v e r s e s e c t i o n showing the o r i e n t a t i o n of the t h i r d and f o u r t h d i v i s i o n s o f t h e c a r p o s p o r a n g i u m . Shown i n s u r f a c e view i n F i g u r e s 9g and 9h. See a l s o F i g u r e 8a.  Figure  9g.  S u r f a c e view o f c a r p o s p o r a n g i u m t h a t h a s u n d e r g o n e t h r e e d i v i s i o n s and c o n t a i n s 4 c a r p o s p o r e s . See a l s o F i g u r e 8a.  Figure  9h.  As  Figure  9i.  Transverse containing  Figure  9j.  S u r f a c e view of a c a r p o s p o r a n g i u m c o n t a i n i n g carpospores (only the top 4 are v i s i b l e ) .  in Figure  Porphyra  9g  f  but  a different  plane  cf  division  focus.  s e c t i o n through a carposporangium 8 carpospores (only 4 v i s i b l e ) .  nereocystis -  8  spermatogenesis  Figure  9k.  S u r f a c e view of t h e f o l i o s e p h a s e s h o w i n g v e g e t a t i v e c e l l s with a s i n g l e s t e l l a t e chloroplast..  Figure  91.  S u r f a c e view o f s p e r m a t a n g i a a few d i v i s i o n s .  which have u n d e r g o n e  82  Porphyra  nereocystis -  spermatogenesis  Figure  10a,  Transverse section spermatangium.  Figure  10b,  S u r f a c e view o f m a t u r e s p e r m a t a n g i a l r e g i o n o f the t h a l l u s . I n d i v i d u a l spermatangia are not clearly delimited.  Figure  10c.  T r a n s v e r s e s e c t i o n t h r o u g h a mature s p e r m a t a n g i u m which i s 8 s p e r m a t i a deep.  Porphyra  nereocystis -  through  an immature  carposporogenesis  Figure  10d.  Transverse s e c t i o n through a v e g e t a t i v e c e l l region of the t h a l l u s . Note t h e s i n g l e s t e l l a t e c h l o r o p l a s t , c e n t r a l p y r e n o i d and l a t e r a l nucleus.  Figure  10e.  Transverse s e c t i o n through a carpogonium. Note i t s s l i g h t l y more e l o n g a t e s h a p e compared t o t h e vegetative c e l l s .  Figure  10f.  T r a n s v e r s e s e c t i o n s h o w i n g two c l o s e l y a p p r e s s e d carpogonia. Note t h e s p e r m a t i u m (arrow) and f e r t i l i z a t i o n canal.  Figure  10g.  T r a n s v e r s e s e c t i o n o f two c a r p o s p o r a n g i a that the f i r s t d i v i s i o n i s p e r i c l i n a l .  Figure  10h.  T r a n s v e r s e s e c t i o n o f two c a r p o s p o r a n g i a . The one on t h e l e f t i s a t t h e s t a g e shown i n s u r f a c e v i e w i n F i g u r e 1 0 i , whereas t h e one on t h e r i g h t i s a t t h e s t a g e shown i n F i g u r e 1 0 j . T h i s d i v i s i o n s e g u e n c e i s t h e same a s t h a t shown f o r P o r p h y r a g a r d n e r i i n F i g u r e 8b.  Figure  10i.  S u r f a c e view carpospores.  Figure  10j.  S u r f a c e view o f a c a r p o s p o r a n g i u m containing 8 carpospores (only the top 4 are v i s i b l e ) . See a l s o F i g u r e 10h.  Figure  10k.  S u r f a c e view o f m a t u r e c a r p o s p o r a n g i a . I ti s d i f f i c u l t t o determine the boundaries o f a s i n g l e carposporangium.  Figure  101.  T r a n s v e r s e s e c t i o n showing two c l o s e l y a p p r e s s e d c a r p o s p o r a n g i a w h i c h have u n d e r g o n e more t h a n one d i v i s i o n i n the plane o f the t h a l l u s .  c f a carposporangium  showing  containing 4  84  Figure  11a.  Porphyra n e r e o c y s t i s * t r a n s v e r s e s e c t i o n through a r e g i o n o f mature c a r p o s p o r a n g i a . I t i s d i f f i c u l t t o determine the o r i g i n a l boundaries o f a s i n g l e carposporangium. Porphyra  Figures  11b  thuretii  -  spermatogenesis  and 11c. S u r f a c e view the spermatangia.  shewing  early divisions  Figure  11d.  T r a n s v e r s e s e c t i o n t h r o u g h an spermatangium.  Figure  11e.  M a t u r e spermatangium with s u r f a c e dimensions b/4 and c o n t a i n i n g a t o t a l o f 64 s p e r m a t i a .  Figure  11f.  T r a n s v e r s e s e c t i o n through a mature spermatangium. Porphyra t h u r e t i i  of  immature a/4,  - carposporogenesis  Figure  11g.  T r a n s v e r s e s e c t i o n through a carpogonium a p p e a r s t o have v e r y s l i g h t b i p o l a r prototrichogynes.  Figure  11h.  T r a n s v e r s e s e c t i o n showing t h a t t h e f i r s t d i v i s i o n o f the carposporangium i s p e r i c l i n a l .  Figure  11i.  T r a n s v e r s e s e c t i o n o f a c a r p o s p o r a n g i u m t h a t has u n d e r g o n e s e c o n d and t h i r d a n t i c l i n a l d i v i s i o n s . A s u r f a c e view i s shown i n F i g u r e 1 1 j .  Figure  11j.  S u r f a c e view of a c a r p o s p o r a n g i u m undergone t h r e e d i v i s i o n s .  Figure  11k.  S u r f a c e view o f a mature c a r p o s p o r a n g i u m c o n t a i n i n g 8 c a r p o s p o r e s {only t h e t o p 4 a r e visible). See F i g u r e 111.  Figure  111.  T r a n s v e r s e s e c t i o n o f a mature  which  which  has  carposporangium.  86  PART I I I - CULTURE STUDIES  Introduction The  first  Janczewski  culture  (1873) and,  demonstrated  that  although  Agardh  P.  £•  l e u c p s t i c t a germinated (1878)  and  however, B e r t h o l d abnormal until  and  they  and  found  (1924), K y l i n and  Rees  Porphyra  anomaly  and  i t  germinate  that  the  standards, (Roth)  J , Agardh)  and  Thuret  filamentous  growths  the c a r p o s p o r e s remained foliose  phase.  o f s e v e r a l Japanese  produced Dangeard  a  and  the  (1939) t h o u g h t  that  from  same  were  unchanged  Yendo  (1919)  Porphyra  species  filamentous  (cited  obtained  suggested  and  are  to give the  reported  that  rhizoid-like the  by  (1882) o b t a i n e d s i m i l a r r e s u l t s ;  i n t o the  also  t h e t h a l l u s and  within  made  phase.  Drew, 1956,  results  with  Grubb p.  574)  European  species.  Kunieda  from  thought  (1922),  (1940)  current  (Lightfoot)  Berthold  carpospores  they  by  were  i n t o a f i l a m e n t o u s phase.  concluded that  the  Porphyra  crude  laciniata  germinated  germinated  of  t h e c a r p o s p o r e s o f Porjojiyra p u r p u r e a  C.  Bornet  (as  studies  foliose  carpospores  that  that  the e n t i r e carpogonium the  carpospores i n the  foliose  phase.  Okamura  carpospores  which f u n c t i o n e d  thallus  growth.  germinated the  directly  They into  carpospores  was  an  i s released  pass  released  immature  that  f i l a m e n t o u s phase  finally  filaments  (1929) s u g g e s t e d  the  the  summer  autumn when t h e y et  germinated t o absorb  a l . (1920) to  nutrients for  stated  that  foliose  thalli,  germinated  produce  mature Ueda  immediately  87 into  small  plants  which  l a s t e d through the  m c n c s p o r e s i n t h e autumn and again.  Kusakabe  Drew the  (1929) s u p p o r t e d  phase r e s u l t i n g  known  to  was  from m a t e r i a l c o l l e c t e d Drew*s various  findings  Porphyra  Krishnamurthy, culture  and  1959;  cytological  environmental  Porphyra  Such  amount o f  data  Kurogi to  on  Porphyra.  the  He the  pioneering effects  of  1953b)  into  the  work w i t h  1964a,  1  a  did  is  net  determine  for  optimal  essential  having  morphological  life  which  light  phase. of  to  of  excellent intensity,  efficient  a  the  great  investigators history 4 species, phase  followed studies  water  of and  formed  conchospores  Kurogi  and  phase.  conchocelis  produced  the  growth  amassed  first  carpospores  These  surprise that  of the  series  photoperiod,  1964c}.;  one  The  1958;  1964b,  was  foliose  (1892)  Hollenberg,  conchocelis  the  was  Batters.  Batters  the  phase.  branches  rjjsea  of  complete  that  Scotland.  i t i s t h e r e f o r e no  biology  germinated  by  1955;  i n t h i s area,  conchocelis  conchosporangial germinated  the  (1953a,  elucidate  obtained  leaders  of  responsible  and  t o p o i n t out  a number of i n v e s t i g a t i o n s of  information  cultivation  Japanese are t h e  coast  studies  parameters  reproduction.  the f i r s t  (Graves,  Ccnway,  thalli  view.  described  stimulated  species  foliose  as C o n c h o c e l i s  first  o f f the  produce  from c a r p o s p o r e g e r m i n a t i o n  phycologists  This s h e l l - b o r i n g alga  produce the  this  (1949, 1954a, 1954b) was  filamentous  already  thereby  summer t o  on  that this the  temperature.  *In some o f t h e J a p a n e s e l i t e r a t u r e t h e c o n c h o s p o r a n g i a l branch is r e f e r r e d t o as a m o n o s p o r a n g i a l b r a n c h and c o n c h o s p o r e s a r e c a l l e d monospores.  88  desiccation  and  conchocelis Hirano, 1959;  salinity  phase  1955a,  and  cn  1961;  (Yamasaki, Migita, and  of  1954a;  Schramm,  Matsudaira,  1958;  of  the  Matsumoto,  1966;  the  1961,  Iwasaki,  or  1962;  1956b;  K u r o g i and  1959^  Ogata,  Sato,  dealt  phase 1S63;  with phase  Kurogi,  Kirita,  and  Sato,  conchocelis  1967;  the  Kurogi,  K u r o g i and  1S71;  foliose  of  (Kurogi  J a p a n e s e work has  1955,  1959;  1971)  1965,  growth  Ogata,  1956a,  al.,  Other  the  1959a; T a n a k a ,  et  maturation  liberation  Hirano,  Akiyama,  K u r o g i e t a l . , 1967). aspects  and  conchospore  Kurogi  1962a, 1962b; K u r o g i and  various  growth  1955b; K u r o g i and  Kurogi,  1967;  on  1956;  1970;  Ogata  (Iwasaki Iwasaki*  and 1965;  Imada e t a l . , 1972). These  studies  manipulate  the l i f e  storing Migita  as  of  nutrient  isolated  the b a c t e r i a l  flora  growth  investigators  i n the s p e c i f i c  by t h e  the e f f e c t branch  them t o d e v e l o p  (Migita,  that  by  few  the  formation  and  such  1964,  1966,  to  novel  nets t o -20°C  has  been t a k e n  foliose  and  1967a;  Japan  c o n d i t i o n s of  and  conchospore  on  release  to  1971,  observe 1974).  also  formation  T s e n g e t ajL*  photoperiod  is  and  have  spore  study  then i n t r o d u c e  thalli,  ( T s u k i d a t e , 1970, outside  t o the  thallus  of the t h a l l i ,  c o n c h o c e l i s phase.  of temperature  species  1975).  approach  affect  effectively  Porphyra  s t r a i n s i n t o axenic c u l t u r e s of  interested release  their  s p o r e l i n g s on  reguirements  the b a c t e r i a A  of  cultivation  Miura,  unigue  the Japanese to  have e n a b l e d  guick-freezing  e t a l . 1971;  characterize  how  and  them f o r l a t e r  Another  enabled  histories  improve c u l t i v a t i o n technigues  have  been and  (1963) s t u d i e d  conchosporangial i n s i x s p e c i e s of  89  Porphyra,  and  investigated branch  the  (1967)  effect  and  of  Rentschler  photoperiod  to  (1961), working  follow  with  have  on c o n c h o s p o r a n g i a l  t h e complete  Porphyra  life  history  defined  c o n d i t i o n s and growth medium.  1961b)  has  done  some  excellent  tenera, in vitro  Koranann  culture  in  life  miniata  and  P. l i n e a r i s  well  European  have  the  using  work on t h e Bird  completing  the  1961a,  Chen e t a l . (1970) and  succeeded  was  (1960,  PojEpJiyra s p e c i e s .  Pophyra  (1967)  f o r m a t i o n by P o r p h y r a t e n e r a .  Iwasaki first  Dring  et  a l . (1972)  histories  respectively,  through  of 5-6  generations. Because thallus  in  of  the  culture,  difficulties there  environmental parameters and  monospore*  importance Kurogi  formation.  suggested  role  in controlling  and  Bird  variety  (1973)  by  found  that and  was d e t e r m i n e d  of  the  (1961)  inducing  pointed  carpospore  temperature  P. l i n e a r i s  photoperiods  out the  formation.  p l a y s an important  Bird  ej£  a l . (1972)  sporulated and  under  suggested  by t h e age o f t h e t h a l l i  rather  a  that than  an e n v i r o n m e n t a l s t i m u l u s . Hy  and  that  studies  f o r spermatium, c a r p c s p o r e  Iwasaki in  few  monospore p r o d u c t i o n .  c f temperatures  sporulation  been  responsible  of photoperiod  (1961)  have  of maintaining the f c l i o s e  culture  P. t h u r e t i i  s t u d i e s of Porphyra  g a r d n e r i . P. r  were i n t e n d e d t o complement  my  nereocystis.  field  studies i n  *In some o f the Japanese l i t e r a t u r e t h e s e s p o r e s which a r e p r o d u c e d by t h e f o l i o s e t h a l l u s and w h i c h g e r m i n a t e t c produce more fcliose thalli, a r e r e f e r r e d t o a s n e u t r a l s p o r e s . The t e r m monospore i n some o f t h e e a r l i e r Japanese literature r e f e r s to conchospores.  90  order to obtain a detailed histories  of  these  picture  three  of  the  algae.  morphological  I  had  the  life  following  o b j e c t i v e s : to i d e n t i f y  the r e p r o d u c t i v e b o d i e s produced  by  foliose  i f different  affect  their  t h a l l u s and germination  see  as has  1970); t o d e t e r m i n e conchocelis  i f  phase  gardneri  i f  so  to  and  branch  the s p e c i f i c  f o r m a t i o n and  factors  characterize  Materials  from used  the  study  site,  m a t e r i a l from  differences  i n c l u d e d : Porphyra  Bock  in  Joe,  Barkley  Beach,  Botanical Porphyra  were c a r r i e d  Preliminary  some s p o r e  as a c h e c k  Porphyra  germination studies for  and  Islet,  monospore  and  other  Cape B e a l e  Sound;  Barkley  and  10°C  and  a  o r W e s t i n g h o u s e F48T12  light Cool  illumination.  of  i n Barkley San  Juan  Island.  Sound.  carpospore  of  for  population  Point,  temperature  used  any  nereocystis: Aguilar  under a p h o t o p e r i o d  were  material  B o t a n i c a l Beach, P o r t Renfrew; P o i n t  out  tubes  with  Execution  were c a r r i e d  Sylvania  out  g a r d n e r i : Second Beach  Beach  Leach  to  Methods  B e a c h , P o r t Renfrew; F r i d a y H a r b o r , thuretii:  g r o w t h ; and  The  Sound;  Cable  general  or type of spore f o r m a t i o n .  Monterey P e n i n s u l a .  Seccnd  And  other s i t e s  sites  the  a  release.  although  i n the t i m e  have  responsible for conchosporangial  conchospore  Most c u l t u r e e x p e r i m e n t s  (fiichardson.  P. t h u r e t i i  f e a t u r e s o f t h e c o n c h o c e l i s phase m o r p h o l o g y and determine  could  been r e p o r t e d f o r B a n g j a P.  and  photoperiods  the  germination studies  12:12  (light:dark),  intensity White The  40  a  of  25-35 f t - c .  W  fluorescent  suitability  of  these  91  c o n d i t i o n s f o r growing previous  culture  b a n g i o p h y c i d a e n a l g a e was  studies  (Hawkes,  determined  u n p u b l i s h e d B.Sc.  in  Honours  thesis). Small t h a l l u s Nalgene  Petri  seawater  medium  control  Marine  plates  diatom  experiments  was  were p l a c e d  (Provasoli,  1971)  and  (Lewin,  t a k e n from and  had  a salinity 1-2  after  development  o f t h e s p o r e s were  Sub-cultures f o r  other  and  Pyrex c u l t u r e these  used  at the  varied  days and,  were  to  fora l l Bamfield  from  30-32%..  i f spore release  was  g e r m i n a t i o n and  observed. experiments  Petri  used  d i s h e s 100x80 mm,  were  p l a t e s and  obtained  culture  in  phase  medium.  450  experiments growth  and  ml were  used  by  number  on  spore  reproduction. in  some  of  experiments.  carpcspores  of  photoperiods  of  15°C.  Both  intensities In  were  and P.  low  enhanced.  nereocystis  of  Porphyra  were  16:T and  (25-35 f t - c ) and  gardneri  germinated temperatures  high  and under  of 7 ° ,  (150-200 f t - c )  10°  light  used.  experiments  host p l a n t  carpospores  8:"l6, 12:12,  Porphyra g a r d n e r i , of  further  conchocelis  Monospores  and  which  enriched  to the standard c u l t u r e c o n d i t i o n s , a  conditions  germination  Seawater system  disposable  (10 ug/ml)  a  p i e c e s were removed and  spores to fresh  addition  GeO  1966).  the seawater  the t h a l l u s  cf  Provasoli's  of  sufficient,  In  100x15 am  50 ml  were examined  transferring  in  with  growth  Station  Cultures  pieces  on  the  cultures  development  of  monospores  were a e r a t e d o r a g i t a t e d  were added  t o some t o  Conchospore  cultures  see of  i f both  growth P.  and  of  pieces  could  gardneri  be and  92  f»  thuretii  light  were m a i n t a i n e d a t  intensity The  and o y s t e r  cn  of  the  conchocelis  i n egg s h e l l ,  shell  For  12:12  o f 200-300 f t - c was used  growth  P. n e r e o c y s t i s  10°C,  clam  of the e f f e c t  aerated.  for these  cultures.  shell  (Saxidomus q i g a n t e u s ) observed.  of temperature  and p h o t o p e r i o d  c o n c h o s p o r a n g i a l branch f o r m a t i o n , the f o l l o w i n g  were Only  used: 10°C  7°,  was u s e d  Temperature which  were  release. table  10°,  at  and  manipulated  in  from  regimes.  8:16"; f r o m  10°C,  16:~8  to  10°C,  12:T2  t o 7°c, 9:15".  10°C  ana  These  20-40%» ,  desiccation,  Cultured  conchocelis  plexiglass culture site  in  the  c u l t u r e tube  to obtain  In  another s e r i e s of  to  was b a s e d  gardneri: 7 ° C , 8:16;  1 3 ° C , 11:'13~  in  phase  an e f f o r t  was  from  subjected  to induce  phase  was  and also  host  plant  placed  a  on t h a t  depth of  o f 7 m. the  to  conchospore  pH o v e r t h e r a n g e 7-10, s a l i n i t y  agitation,  and from  Porphyra t h u r e t i i :  conchocelis  at  temperature  and  t u b e and p u t o u t i n t h e f i e l d  autumn  gradient  i n c l u d e d : Porphyra  15°C, 16:8  factors  conchospore  temperature  to different  1 0 ° C , 8:16";  included:  main  t o 10°C, 10: 14* and 7 ° C , 9:15;  conditions  release.  the  7 ° C , 12:TT  12:12  flgreocystis  other  These  from  10°C,  7°C  a  7°-20°C.  to  Porjghi£a n e r e o c y s t i s :  several  attempt  were t r a n s f e r r e d  10°C, 12:12  Porphyra  an  also  were m a i n t a i n e d on  temperatures  photoperiod  from  thuretii.  and p h o t o p e r i o d were  experiments c u l t u r e s  conditions  15°C and 8:16, 12:12, 16:8", 25-35 f t - c .  f o r P.  Cultures  A  phase o f P. g a r d n e r i and  { C r a s s o s t r e a g i q a s ) was  studies  and  from  exudate.  in  a  special  at  the  study  The d e s i g n o f t h e  *Biomonitor*  which  is  93  manufactured  by t h e  (Schlichting,  BioCo.nt.rcl Company, P o r t  1975a, 1S75b,  Sanilic,  Michigan  1976).  Besults  *•  Porphyra  1.  gardneri  Monospores  Soon a f t e r in  February  granular  the  of to  cells  the  The  which  release  cell  (Figure  of t h i s r h i z o i d  a  long  in  the  to  thallus  ( F i g u r e s 12e but  growth c e a s e d  'host*  formed  4-5  angles  initially,  the  was  the  field a  fine  stellate  best developed  Following  entire  distal  which  the  in  appearance  margins  began  were 25-30 urn i n  cell  rhizoid an  stage  when t h e y and  they  12f). reached  1-2  (Figure  to ensure  tissue before  that  elongation,  divisions the  days 12b). the  the f o l i o s e a  uniseriate thallus  i n i t i a l ones t o form and  after  protuberance  i s probably  Following  divisions at  obscured  monospores  rhizoidai  the  commences g r o w t h .  12d).  the  the  zone developed  monospores g e r m i n a t e d  function  and  margin.  in  12a).  out  parallel  the  of  sending  anchored  marginal  appearance  contents,  thousands  appeared  partially  granular  In c u l t u r e the  is  i n the  which were on  granular  diameter  gardneri f i r s t  cells  appearance  chloroplast. those  Porphyra  The spore  thallus  series  of  (Figures  12c  occurred  foliose,  by  at  right  monostromatic  S p o r e l i n g s grew w e l l i n c u l t u r e t h e s i z e shown  eventually died.  in  Figure  12f  94  This increase spring their  prolific i n P.  lateral  During  margin  and  margins  production presence  production  May,  thalli  s p e r m a t i a and  were  common.  a  few  bipolar  monospores were s t i l l  In  i n the  sporelings  being produced.  monospore g e r m i n a t i o n .  phase;  was  found  examined. that  (type l o c a l i t y ) The  indicated  be  at  16:8  the  failure  Culture  number at  by  work  done  most o f t h e t h a l l i  in July  1976,  of sporelings  growth, 3,6% Using  tested  Parker  (v/v)  for  this  solution  one  to reach  the  (1966)  reported  Microcystis  concentration  o f d(-)  in  the  other  a t Hopkins  Marine  as  m a n n i t o l was  of  15  a  a mannitol  monospcric.  however,  they  plant.  in this on  In  group  sporeling  c o n c e n t r a t i o n of  ( L i n n a e u s ) C.  guideline  tested;  Point  laminarialean  mannitol  pyrifera  at  'host*  major p h o t o s y n t h a t e s effect  monospores  maturity suggested  f o r the  the  8:16.  were e x c l u s i v e l y  reguirement  of the  temperatures  collected  o f t h e o c c u r r e n c e o f P o r p h y r a g a r d n e r i on because  good  or  plants  view  mannitol, I  some  for  of  12:12  have a p h y s i o l o g i c a l  is  that  essential  may  a l g a e and  the  g e r m i n a t i o n t o produce  c o n s i d e r a b l y lower than  Station confirmed Joe  spore  I n t h e autumn, moncspore  to  Monospores were b e i n g p r o d u c e d populations  July,  Under a l l p h o t o p e r i o d s and  however,  g e r m i n a t i n g was  their  b u t n o t t o t h e same e x t e n t  monospores underwent b i p o l a r  foliose  and  along  spring.  culture aeration  tested,  the  carposporic, although  p r o d u c t i o n a g a i n became p r e d o m i n a n t , as i t was  June  great  monospores a l o n g  carpospores  By  the  throughout  releasing  became a l m o s t e x c l u s i v e l y of  explains  g a r d n e r i abundance i n t h e f i e l d  months. distal  monspore  a  1,8$ no  Agardh. and  0.9%  enhancement  95  of  growth In  occurred. another  were added  to  Laminaria  resulting  2.  the  was  host tissue  experiment monospore  germinated  Spermatangium  in  And  the  was if  response,  monosporic  kept  1 containers at  Spermatia  formed  is  under  Carpospore  They  spores  carpospores germinated 4-7  um  wide.  parietal  were  both  the  i n 2-3 The  14-20  12:12  sere  that  released  *host*  16:"8~,  plant  were  300-400 f t - c .  U n f o r t u n a t e l y the  C o n c h o c e l i s Phase  been f o r m e d by  divisions is  produced  not  diameter  occurred known.  the  conchocelis  carpospores  Released and were  contained  Under a l l t e m p e r a t u r e s germinated  or  after  ( F i g u r e 12g)  phase  by  one  give conchocelis filaments that  of  To  photcsynthetic  c a r p o s p o r e s were  ( F i g u r e 12h).  tested,  and  carpogonium  formed.  - The  in  the  formation.  or  whether t h e y had  um  but  suggested  their  photoperiods.  first  days t o cells  I)  attached to the  10°C,  been  chloroplasts  photoperiods  without  Formation  inducing  Whether s u b s e g u e n t had  control,  photoperiodic  Germination  were v i a b l e  more d i v i s i o n s . the  a  in  thalli  At t h e s t u d y s i t e May.  control  c f s p e r m a t a n g i u m and  died before carpospores  3.  the  (discussed i n Part  determine  i n 20  A  thallus  maturity.  Carpogonium  important  this  in  grow t o  time  with l a t i t u d e  photoperiod  cultures.  before those  s p o r e l i n g s d i d not  formation  p i e c e s of L a m i n a r i a  2  a l s o r u n . , Monospores i n the c u l t u r e s c o n t a i n i n g  Variation  thalli  1 cm  into  and the  96  conchocelis ether  phase.  C a r p o s p o r e s were p r o d u c e d  populations  carpospores  produced  low  compared t o  and  germinated i n t o The  which by t h e  fragmentation  colonies.  of  i n some c a s e s  functioning  as  were i n i t i a l l y long  and  release  s h o r t and  branched  Aeration  to  was  stubby  (Figure (Figure  i n diameter  germination  was  and  produce found  to  1 2 i ) , but  locality  was  were v i a b l e  soon  no  formed  filaments  by  swellings  evidence  o f them  seen.  (Figure  12j).  13a).  branches formed. 13a),  By  24  2-3  the  foliose  but  later  weeks  Released  after  be  of  spore-like  weeks c o n c h o s p o r a n g i a l  occurred  22-30 um  Single  (Figure  monosporangia  10-12  Propagation  readily.  the  phase. and  occurred  cf  number  they  phase grew p r o l i f i c a l l y  were formed  After  type  but  conchocelis  a l l  The  a t the  populations,  the t y p i c a l  tuft-like  examined.  population  more n o r t h e r n  conchocelis  numerous  were  by  became  conchospore  conchospores  days they  for  were  underwent b i p o l a r  phase  essential  They  (Figure  good  13b).  conchospore  germination. When  grown  characteristic Growth i n t h e in  the  clam  conchocelis The darkness  in  pinnately eggshell  and  ability  conchocelis  phase  morphology  In  to the  older  conchocelis After  8  were p a l e p i n k ,  to the  light.  but  of  a 13c).  prolific  growth  portions  phase t o  months  had  (Figure  b r a n c h e s became i r r e g u l a r l y  tested.  w e l l when r e t u r n e d  conchocelis  poor compared  shells.  of the  filaments  the  branched  was  oyster  phase t h e  was  shell,  of  the  lobed.  survive i n  total  darkness  the  were a l i v e  and  grew  97  4.  ;  C c n c h o s p o r a n g i a l Branch  The  effects  of  ccnchosporangial  Formation  photoperiod  branch  and  temperature  formation  were  C o n c h o s p o r a n g i a l b r a n c h e s were i r r e g u l a r l y  branched  in  making  size,  assessment Dring  length  and  of t h e i r  (1967)  considered  on  Porphyra  it  does  conchosporangia. to obtain  culture was  the  10°C,  12:12. under  One lcng  formed  The  presence  o r two  i t  was  (1970)  has  number  of  encountered i n In  each  o f abundance made.  a t 16:8,  at  8:16,  b u t most  and  cultures  Conchosporangial branches  but b e c a u s e  possible  formed  of  culture  chamber  to determine p r e c i s e l y  the  f o r c o n c h o s p o r a n g i a l branch f o r m a t i o n .  The 10°C  7°C.  Conchospore  formation,  phase  f o r c c n c h o s p o r a n g i a l b r a n c h e s t o form s o o n e r a t  Although  proved  assessment  vegetative.  not  actual  by  of conchosporangial branches  branches formed  was  guantitative  insurmountable.  ccnchosporangial tranches  15°C t h a n a t  5.  proved  at a l l 3 temperatures,  was  the  varied  t e c h n i g u e used  difficulties  or absence  and  Richardson  determine  data  The  examined,  conchocelis as  technical  this  optimal temperature  and  not  days remained  malfunctions  trend  tenera  because,  n o t e d and a g u a l i t a t i v e at  density,  abundance d i f f i c u l t ,  impractical  pointed out,  trying  branch  on  i t  getting  impossible.  m a i n t a i n e d a t 7°C,  Release was  easy  them t o  t o induce c c n c h o s p o r a n g i a l branch release  Conchospore 10°C  and  8:16,  conchospores release occurred  12:T2,  consistently in cultures  w i t h t h e most  prolific  98  release  occurring  ccnchcspore  10°C,  £.  12:T2  at  the  r e l e a s e was  latter  induced  photoperiod.  by t r a n s f e r r i n g  A  small  a culture  from  t o 7°C, 12:T2.  Porphyra n e r e o c y s t i s  1.  Carpospore  No was  evidence  Germination  of  mcncspore  - The  C o n c h o c e l i s Phase  p r o d u c t i o n by t h e  foliose  thallus  found. Carpospores  Porphyra  nereocystis  released  from  degrees  days they  produced which  germinated  chloroplasts. carpospores production  had  c/4,  i n diameter  the  into  o f monospores by t h e  although i t readily  regenerated  varying  Part I I ) .  Released After  filaments  which c o n t a i n e d and  the  photoperiods  conchocelis  fragments  of  2-3  4-6  um  parietal tested,  phase.  c o n c h o c e l i s phase was from  were  undergone  ( F i g u r e 13d).  of  Onder a l l t e m p e r a t u r e s germinated  see  of  Carpospores  conchocelis  cells  populations  examined.  and  t o produce  ( F i g u r e 13e),  a l l  which  ( b o t h c/2  were 8.0-15 um  by  were  carposporangia  of d i v i s i o n  carpospores  across  were  No  observed, vegetative  filaments. A and  number  of  irregular  s w e l l i n g s formed a t b o t h t h e  apex o f f i l a m e n t s ( F i g u r e s 13f and  11-15  um  13-17  um  mistaken  across across  ( F i g u r e 13h)  ( F i q u r e 13i)  and  also  f o r c c n c h o s p o r a n q i a l branches.  specific  function  such  as  Beaded  square-celled  formed  formed under a l l p h o t o p e r i o d reqimes any  13g).  and  were  These two  tested.  base  branches branches initially  branch  Whether t h e y  types have  the p r o d u c t i o n c f monospores i s  99  not  known.  T h e y a p p e a r t o form  filaments to ccnchosporangial Conchosporangial were t y p i c a l l y branches  were  and  cluster  uniseriate,  The cells  thick  pinnate  small  orientation  To t e s t  admitted  on  removed to  cell If a  from  grow,  the  producing filaments.  branches  were two  .  the l i g h t  conchocelis which  14d).  filaments  Conchosporangial  this,  s u r f a c e . , The  suggested  that  they  a c u l t u r e d i s h was c o v e r e d  grew  Under t h i s  toward  vertical were so t h a t  treatment a l l  the l i g h t ,  indicating  darkness,  vegetative  phototropic.  in  barnacle  of the  14b).  conchocelis  the s h e l l  branches  branches  filaments  was  (Figure  was  continue  a t one end o n l y .  conchocelis  It  cells  (Figure  A t t h e end o f 8 months i n  barnacles  The  f o r m e d i n t h e s h e l l a n d t h e n grew c u t c f i t  are p o s i t i v e l y  The  Each c e l l  branches or vegetative  branching  these  ccnchosporangial they  and 14a).  chloroplast, thick  the vegetative  colonies  of  phctotropic. was  10-15 weeks and  (Figure 14c).  branches i n i t i a l l y form  13k  p o r t i o n s o f some c o n c h o s p o r a n g i a l  developed  light  would  vegetative  13j).,  um a c r o s s .  branches  i t  When grown i n s h e l l  to  (Figures  between a d j a c e n t  mere c o n c h o s p o r a n g i a l  older  (Figure  b r a n c h had a s t e l l a t e  filaments  from  appeared a f t e r  16-25  conchosporangial  vegetative either  branches  branches  p i t plugs  of  transition  candelabra-shaped  conchosporangial wall  a  complete  were as d e e p l y  phase was  observed  were on t h e s t i p e  pigmented as those  i n the f i e l d  grown  growing i n  o f an o l d N e r e o c y s t i s  plant.  t a k e n back t o t h e l a b o r a t o r y and t h e s u r f a c e l a y e r s o f d i s s o l v e d by p l a c i n g them i n a 1% s o l u t i o n o f  Na EDTA 2  100  for  12 h r  patches to  (Prud'homme van of  conchocelis  culture dishes.  square-celled Porphyra  phase  suggests that  2.  This isolate  and  The  and  its  of branch  conchosporangial  branches  tested.,  They At  first  8:16  to  this  co.nc.hospores  Formation  formed  branches  of  of  nereocystis.  and were  under  photoperiod examined. a l l  i n cultures at  3  which  formed  At  and  formed  a t 15°C  on 10°C,  photoperiods  15°C  most c o n c h o s p o r a n g i a l b r a n c h e s  Conchosporangial  beaded,  typical  release  to Porphyra  formation  appeared  formed  morphology  temperature  conchosporangial  12:12.  failure  Small  transferred  branches  general  i t i s referrable  effects  Hoek, 1966).  grew w e l l and  conchosporangial  C o n c h o s p o r a n g i a l Branch  The  van den  f i l a m e n t s were removed and  nereocystis.  conchocelis  S e i n e and  16:8, at  or  15°C.  were s h o r t  and  stubby. ,  3.  Conchospore  None  of  succeeded.  the  attempts  Because  autumn i t was seawater  the  Release  to  suspected  that  were  resulted The  decreasing factors;  possibility caused  release  i n the  late  daylength  and  however, ncne o f  higher temperatures  c o o l e r t e m p e r a t u r e s and  i n conchospore  Nereocystis  the  critical  t r a n s f e r experiments, from to  conchospore  porphyra n e r e o c y s t i s appeared  temperature  photoperiods  induce  and  longer  shorter photoperiods,  release.  that  some  conchospore  chemical release  cue was  from  the  aging  investigated  by  101  placing  pieces  conchocelis The found  Neither  conchocelis  suggesting  of  zone  exudate, i n  t r e a t m e n t s was  (Drew and  No  with  Richards, be  branches subjected  1,  to  were  2 and  has  1953;  been  Mumford,  a factor  ccnchospores  the  successful.  some Porph y r a s p e c i e s  that desiccation could  release.  ccnchosporangial  or s t i p e  of these  phase  intertidal  conchospore  at  mature s t i p e ,  phase.  i n the  19 73a)  of  affecting  released  3 hr  of  by  drying  20°C. Because  the  substrate  i t was  reguired  for  ranging The  from  other  conchocelis suspected  conchospore pH  7-10  that  phase  grows  there  may  be  Six  cultures  release.  were t e s t e d .  cultures  survived  but  At pH the  in  a a  10 t h e  pH  calcareous specific with  culture  pH a  pH  died.  g u i c k l y changed t o  pH  8-9. Changes  in  salinity  had  no  effect  in  triggering  conchospore r e l e a s e . The  conchosporangial  c u l t u r e tube a l s o f a i l e d  C.  Porphyra  1. No °f  evidence  Released  of  monospore w  a  s  field  in  the  and  production  were  germinated  phase  across  i n general  Conchocelis by  the  Phase foliose  thallus  found.  carpospores  conchocelis and  i n the  produce ccnchospores.  C a r p o s p o r e G e r m i n a t i o n - The  14e)  out  thuretii  Porphyra t h u r e t i i  {Figure  to  branches put  {Figure  14f).  15-22 after The  morphology and  um 2-3  in days  filaments c o l o u r they  diameter to  give  a  were  4-5  um  were  distinct  102  from t h o s e o f P o r p h y r a the  vegetative  which were  filaments  in  my  formed  studies  Herbarium  were made from phase  and P. n e r e o c y s t i s . long,  fine,  In culture  hair-like  'tufts*  purple-gray.  Early identify.  gardneri  was  this  so  I collected  thalli  which I c o u l d  s p e c i m e n s and c o n c h o c e l i s material  distinct  f o r future  that  phase  study.  I was l a t e r  The  able  not  cultures  conchocelis  to refer  i t to  Porphyra t h u r e t i i . The  vegetative  branches  (Figure  branches formed in  older  (Figure In  one  14h).  a  formed followed were  the  by  released  germination (Figure as  i t  initial  a  15e). did  in  rhizoid  long The  produce a b a s a l  12  but  colonies  weeks,  abundant  conchosporangial  branches  release  exhibited  (Figures  ( F i g u r e 14k).  are  not  pronounced  known. amoeboid  (14-17.5 um i n d i a m e t e r ) and  within  rhizoid  15a-d).  formation  this  rhizoidal  (Figures  short,  Following  elongation,  pad  were  processes  occurred.  (Figure  conchosporangial  at t h e time o f t h e i r  of these  In  the i n i t i a l  protuberance  stage of  was  formed  d i d n o t d e v e l o p t o t h e same  Porphyra g a r d n e r i  protuberances formed  of  conchospore  up  they  beaded  phototropic.  the  conchospores  germination  weeks  and  candelabra-shaped  end  not observed  so the d e t a i l s  days  long  within  c o n c h o s p o r e s rounded few  8-14  were p o s i t i v e l y at  irregular  Initially  they formed  c h a n g e s o f s h a p e and movement the  developed  After  experiment  14 j)  release,  Freshly  (Figure  14i) which  Conchospores or  14g) .  cultures  conchospores (Figure  filaments  a  sporelings., few  15f) and  short,  Following broad  partially  15g, 1 5 i , 15j) .  extent the  rhizoidal  coalesced  to  103  Growth parallel  15g).  cell  divisions  (Figures  which  resulted  in  a  and  effect  formation  was  at  then  8:16  entirely  at  In  but  most  of  thallus  to about  (Figure  culture, t h a l l i  15h)  and  thallus  there  grew t o  was  about  on  conchosporangial  branch  a t 8:16  branches.  first  consisted  In  1  almost  out  cf  3  conchosporangial branches forned  cultures  a  Formation  Cultures  of  the  perpendicular  C o n c h o s p o r a n g i a l branches formed  12:12,  couple  series  filament  ceased,  conchosporangial a  uniseriate  t h e s e young s p o r e l i n g s  photoperiod  studied,  of  experiments 16:8,  In  growth  of  with a s e r i e s of  a parenchymatous, monostromatic  C c n c h o s p o r a n g i a l Branch  The  an  a  biseriate  15j).  before  in  time  a p i c a l growth zone.  i n length  2.  to  began  remained u n i s e r i a t e o n l y  at  gave r i s e  thallus  resulted  Sporelings  15i  definite am  foliose  stage,  eventually  1  the  d i v i s i o n s which  (Figure 4-7  of  remained  vegetative  at  at  this  photoperiod.  3.  Conchospore  Conchospore at  10°C,  7°C,  9:15.  conditions other  12:1~2,  Release  release and  by  Subseguent failed  to  occurred  under two  transferring exposure  of  conditions  from  other  10°C,  12:12  to  cultures to  these  induce conchospore r e l e a s e  f a c t o r s were a l s o  involved.  tested:  i n d i c a t i n g that  104  Discussion  1.  Monospores  Porphyra the  gardneri  Northeast  Pacific  monospore  cycle.  monospore  production  British  Columbia  observed i n . form  Although by  they  This  P o r p h y r a which  small,  1974).  P.  is similar  gardneri  which  produce  (Kurogi,  Why  In  and  environmental species  provide  This  implies  related  to  kuniedai  both  play  a role that  monospore  this  was  thuretii  do  not  (Kurogi,  and to  70  of  1972;  production P.  yezcensis  mm  in  length  Shinmura  (Shinmura, be so  in  which  1S74).  prevalent  which  monospore-forming  monospore  of  the  A  the  study  in  guestion.  the  production  phenomenon., that  monospore  predominant i n  that  up  triggering  into this  in  monospore  should  under  plants  10 J a p a n e s e s p e c i e s  a l l seasons  demonstrated  (1961) s t a t e d  to  thalli  intriguing  reported  a p p l i e s t o most s p e c i e s  taneqashimensis  factors  g a r d n e r i was  t e m p e r a t u r e s may Kurcgi  the  insight  studies  Porphyra  P.  from  predominant  species  Porphyra  prolific  production  conditions  grow and  My  on  a  intertidal  p r o d u c e monospores  tc Porphyra  known  a l . (1975)  high  think  its  to  J a p a n e s e s p e c i e s i s an  et  such  s p e c i f y which  I  monospores t h r o u g h monospore  has  contrast  monospores  1972),  liberates  in  known t o  Shirmura,  which  n e r e o c y s t i s and  is  are  Porphyra s p e c i e s  Conway  d i d not  Porphyra  area.  only  Ocean  monospores, a s i t u a t i o n  this  may  i s the  the  shorter  production  spring  and  photoperiods  i n inducing  monospore  autumn.  and  by  the  fcliose  lower  formation,  water t e m p e r a t u r e seems t o be  liberation  by  closely  thalli  of  1.05  LPJphyya  k u n i e d a i and  Eangia,  Sommerfeld  P.  tenera.  and  Nichols  In  culture  experiments  with  (1973) d e m o n s t r a t e d t h a t  water  t e m p e r a t u r e a f f e c t e d monospore p r o d u c t i o n . indicated  that  there  is  a correlation  moDCspere p r o d u c t i o n  by I.  I g z o e j a s i g and  species,  thallus  size  determines  produced.  I n Porphyra  than the  2 mm field The  the  high  do  a  not  form  production  unigue  P. , k u n j e d a i  of  for it of  be  are  thalli  more  and  carpospores  early  (1961)  on  summer  is  reported t h i s  for  yezoensis.  one  of the  cycle,  economic  Porphyra g a r d n e r i  most v a l u a b l e s p e c i e s on a s s u m i n g t h a t a method  a s y n t h e t i c s u b s t r a t e can  major  some  1961) .  Kurogi  commercial c u l t i v a t i o n , on  In  and  monospores  example,  g a r d n e r i i n the  B e c a u s e o f i t s monospore potentially  £. tej$e.r_§.  monospores, s p e r m a t i a  phenomenon. P.  also  monospores e v e n when c o n d i t i o n s i n  i n Porphyra  and  (1961)  between n u t r i e n t s  whether  Deda f o r  are s u i t a b l e (Kurogi,  same t h a l l u s  not  angusta  Kurogi  be  importance  developed.  could  our  of  growing  Japanese  have a monospore c y c l e  coast  species (Kurogi,  1972).  2.  Spermatangium  My  initial  carpogonium suggested The  that  (less  daylength), a  field  Carpogonium  photoperiod  in was  of spermatangia than  Formation  observations  formation  formation  culture  And  of  Por.phy.ra g a r d n e r i  at  a  photoperiod  what a p p e a r e d t o be  effect  (see  responsible for their  i n d i c a t e s t h a t spermatangium  photosynthetic  spermatangium  rather  than  the  of  truly  is  I)  formation.  critical  formation a  Part  and  12:12  in  inducing probably  photoperiodic  106  response  as  defined  experimental determine form  and  why some c e l l s  than  Kuroqi  however,  spermatangia  spermatangia  (1961)  certainly  and S u t o  (1965) r e p o r t e d t h a t  12 h r l i g h t ,  carpogonium  produce  Fujiyama  tried  concluded  whereas  from  to  correlate  this  be  Ogata,  an  1975)  was g r e a t e r  formed. spermatanqiuni  with was  others  to  when t h e d a y l e n q t h  and c a r p o q o n i a  that  Critical  seems  {cited  f o r m a t i o n i n P. k u n i e d a i  he  (1970).  t o answer t h i s q u e s t i o n and t o  Photoperiod  factor.  Iwasaki  Bichardson  s t u d i e s a r e needed  carpogonia.  important  by  water not  and  temperature;  the  only  factor  inducing sexual reproduction. When e x p e r i m e n t a l t e c h n i g u e permit thalli  manipulation i n culture,  begin.,  It  selection Such  will  a  new  then  may  (Dr. T. F. Mumford,  Effect  sufficiently  refined  to  of the r e p r o d u c t i v e s t a t u s o f the f o l i o s e era  in  Porphyra  cultivation  be p o s s i b l e t o a t t e m p t  t o improve the c u r r e n t l y  studies  3.  is  already  h y b r i d i z a t i o n and  cultivated  be  will  Porphyraspecies.  going  cn  in  Japan  p e r s o n a l communication) .  Of  Photoperiod  On  Monospore  And  Carpospore  Germination Germination carpospores  of  under d i f f e r e n t spore  type  photoperiod. al.  of  P. g a r d n e r i , photoperiods  i s  induced,  of  Porphyra  P. n e r e o c y s t i s showed i t s fate  that  (1973a) on P o r p h y r a .  In c o n t r a s t , B i c h a r d s o n  gardneri-  and  once  cannot  T h i s i s i n agreement with the  (1972) and Mumford  Bangia.  monospores  P.  a  thuretii  particular  be  work  and  changed of  and C o l e  (1970) r e p o r t e d t h a t  Bird  by et  (1972a) on i n Bangia  107  released  carpospores  cultured  underwent  under a p h o t o p e r i o d  unipolar  germination  cultured  under  a  Conway  (1964b,  less  give  photoperiod 1966)  conditions carpospores spcrelings  to  of  instead  has  bipolar  tested  are  (1977b) have b o t h  4.  of Porphyra  is  the  histories  establishing the  1972)  that  Porphyra  Porphyra lack  when  12 h r  that  that  under  certain  info  bipolar  I n such  all  Kurogi  light.  of  cases I  the  spores  (1961) and Hawkes  monospores and c a r p o s p o r e s  can  be  first of  they  species  r e p o r t o f a c o n c h o c e l i s phase i n t h e  Porphyra  branch  branches  gardneri  have h e t e r o m o r p h i c  and  P. t h u r e t i i *.  life  histories.  which h a v e been c u l t u r e d , (Conway, 1973; Conway  subtumens J . Agardh ex L a i n g  2  only et  Cf  Porphyra a l . 1975)  (Conway and M y l i e ,  a c o n c h o c e l i s phase.  Irregular and  phase  thallus.  s a n j u a n e n s i s Krishnamurthy and  than  can germinate  o f t h e same t y p e . that  and  The C o n c h o c e l i s Phase  This life  more  when  light,  conchocelis  o f the c o n c h o c e l i s phase.  noted  p r e s e n t on t h e same  12 h r  suggested  t h i n k t h a t one s h o u l d be v e r y c a r e f u l being  than  the with  germination  swellings of  types  have been  vegetative  which s u p e r f i c i a l l y r e p o r t e d by s e v e r a l  conchocelis  resemble workers  filaments  conchosporangial (Iwasaki,  »The r e p o r t by Conway e t a l . (1975) t h a t P o r p h y r a a c o n c h o c e l i s p h a s e was b a s e d on my work.  thuretii  1961;  has  It should be n o t e d t h a t i f t h e mode o f s p e r m a t i u m f o r m a t i o n r e p o r t e d by Conway and Sylie (1972) i s correct, then this s p e c i e s does n o t b e l o n g i n t h e genus P o r p h y r a . 2  108  Iwasaki J977).  and  Matsudaira,  Conway and  ncncspores. Porphyra  released  Matsudaira  produce  i t  phase  are  monospores.  by f r a g m e n t a t i o n r a t h e r  Chytrid  infection  spore-like (1919)  by  Migita,  by  Mumford  (Arasaki, 1973,)  when under  branches  which  that  they  were  the never  propagation in  my  monospore  of  cultures  production.  (1973a) i n h i s c u l t u r e s . phase  1960, which  that  conclude  filaments  than  Cole,  produced  maintained  sguarish  of the c o n c h o c e l i s  swellings  work;  conchocelis  observed  found  Vegetative  occurred  also  (1963)  monosporangia because  nereocystis  was  these  definitely  Porphyra  This  that  was  formed  I cannot  I observed  to  and  reported  conchocelis  monospores.  Conway, 1964b; Conway and  (1977)  illumination  swellings seen  Iwasaki  tenera  continuous  Cole  1963;  can  also  produce  to  lendo's  referring could  be  mistaken  for  monosporangia. Under Porphyra  certain  conditions,  nereocystis  monospores, some r o l e  or  they  and may  Porphyra  branches  thuretii  produced  may  these  beaded  (1968),  in  process. branches  as  Krishnamurthy as b e i n g  by  produce  have o t h e r f u n c t i o n s s u c h  i n the s h e l l - b o r i n g  misinterpreted  t h e beaded  playing (1969a)  conchosporangial  branches. Fukuhara species species into  of  Hokkaido,  his  noted  types;  one  in  ccnchosporangial branches which t h e y  which  branches  Porphyra  several  Porphyra  could  be  divided  t h e v e g e t a t i v e f i l a m e n t s and  are a d i f f e r e n t  a r e t h e same c o l o u r .  ccnchosporangial  of  t h a t t h e c o n c h o c e l i s phase of t h e  b e l o n g i n g t o t h e subgenus  two  studies  colour  and  one  in  In the 3 s p e c i e s I s t u d i e d  the  were more d e e p l y p i g m e n t e d  than  the  109  vegetative The  f i l a m e n t s but pinnately  were n o t  branched  f i l a m e n t s grown i n s h e l l how  the  (1959;  shell  1961)  vertical  morphology  was  striking  matrix causes  presented  growth  markedly  p a t t e r n o f the  the  morphology  was  found  that  the  degree  regularity  branching Kornmann, Kim, that  pinnately  branched  showed  (1975)  also  physiological this  shell  that  study  as  . they  as  are a c t u a l l y  reported that vegetative  by t h e  the  cf branching  varied  s p e c i e s develop 1953b;  Migita  and  type of s h e l l .  and  Kim  whereas  under  filaments positive  and  with  1954a;  Migita  (1970)  and  found the  unidirectional conchosporangial  phototropism.  a  positive  phototropism  of  Pcrphyra  brumalis  needed  I  pinnate  Drew,  1962a, 1962b, 1967;  to elucidate  to the branches  'fertile-cell  'plantlets*.  i n agreement  of  Mumford for  the  Mumford.  A  t h e mechanism  of  response.  r e f e r r e d t o as t h e are  is  (1954a) r e f e r r e d  matrix  surface  branches  photographs  Ogata  c o n c h o c e l i s f i l a m e n t s developed  definite  reported  phototropic Drew  light  vegetative a  conchosporangial  Sato,  morphology,  the  branches  and  conchocelis  p c s e s t h e g u e s t i o n of  1953a,  1973a, 1 9 7 5 ) .  diffuse  both  (Kurogi,  Kurogi  Mumford,  under  light  shell  1960;  1970;  affected  S e v e r a l other Porphyra  in  the  c o n c h o c e l i s phase i n s h e l l  that  type.  and  fascinating  noted  the s h e l l  of  i n cclour.  t h i s m o r p h o l o g i c a l change.  some  and  different  Migita one  rows'  she  and  (1961, 1962)  continuous  t h o s e on has  structure  c o n c h o s p o r a n g i a l branches. with M i g i t a ' s c o n c l u s i o n .  the c o n c h o s p o r a n g i a l  observed  branches  in  the  the shell  pointed and  My Migita could  out  should  be  observations (1962) revert  f i l a m e n t g r o w t h which c o u l d r e - p e n e t r a t e t h e  also to  shell.  110  I  a l s o observed Ogata  the  this  (1961) u s e d  internal  speculated  pfi  the  ability  of  i n was  The  was  the c o n c h o c e l i s  the  Rosenvinge,  1975).  field  1931;  at t h e  provided  by  responsible  actual  for  prolonged i s needed  into adaptations  light  and  on  in liguid able  the the  chloroplast  Ogata  growing  of  reported  photosynthesis  leucosticta.  was  He  physiological  phase t o s u r v i v e  low  in  the  (1961,  1971)  even  after  paraffin.  He  t o use  shell  been o b s e r v e d  number  (reported  as  of  Richards,  the  crustose  times  19 53;  site  the  the  concluded matrix  van  report  of  substrate, corallines  f o r the c o n c h o c e l i s  barnacles  algae.  In  1892;  den  Hoek,  1958;  Mumford,  1S73a,  view o f  substrate Drew  Lithothamnion laevigatum being  a thorough survey are  ruled  phase.  out  or  (Batters,  predominant c a l c a r e o u s  L i t hothamnion l a e v i g a t a ) phase  in  M i g i t a , 1959b; B i r d , 1973;  crustose c o r a l l i n e  (1953)  substrate  a  Drew and  study  Richards  before  phase has  I t o , 1959;  conchocelis  5-6.  source.  in  M i u r a and  was  pH  that  elucidated.  continued  embedded  conchocelis  shells  and  pigmentation  shell  a carbon  was  a l . (1977) have r e c e n t l y  conchocelis  from t h i s t h a t  the  n e e d s t o be  phase of Porphyra  i t was  determine  i n which p h y s i o l o g i c a l s t u d y  observed that  as  however,  darkness  ultrastructure, conchocelis  acid production  interestng insight  Sheath et  to  filaments  conchocelis  area  provide  conchocelis. effects  mild  process  darkness i s another should  conchocelis  of the  nereocystis.  sensitive stains  shell;  mechanism o f t h i s  and  pH  of  that t h i s  dissolving  The  phenomenon i n P.  as  a  Fcslie suitable  should a  and  be  made  potential  111  5.  Conchosporangial  Conchosporangial l21Bfel£§ t h u g e t i i 12  hr l i g h t  photoperiod  Branches  branch  was  f o r m a t i o n by  most  or  less,  was  observed.  abundant  whereas The  Porphyra  species  importance  to conchosporangial  and  Bentschler  in  P.  that branch  both  £• t h u r e t i i and  are s i m i l a r  Sasaki  (1972),  under a l l p h o t o p e r i o d s and  abundant  is  involved.  The that  al.  data o f Japanese  1975)  The  Iwasaki,  1962;  i s that  but  of  ether major  Dring  (1967)  Porphvra  tenera.  was  a  short-day,  w i t h P.  gardneri  (1967)  and  were r a r e  under  and  Iwasaki formed  long  days  I t i s n o t known i f p h y t o c h r c m e  (1970),  however,  producion  by  reported  pangja  main c o n c l u s i o n 1961;  of  K u r o g i and  1965;  was  an  Iwasaki  that 'all  or  Sato,  branch  hypothesis  i n conchosporangial several  and  the  workers  1962a,  per 24  hr.  i n t h e autumn and  most J a p a n e s e P o r p h y r a species,  S a s a k i , 1972;  et  Chihara,  formation i n the  Japanese  In the f i e l d winter  and  (Kurogi,  and  subjected  such c o n d i t i o n s  when t h e f o l i o s e  s p e c i e s appears. Kurogi  branch  1962b; K u r o g i  s p e c i e s i s most a b u n d a n t i n c u l t u r e s  12 h r o r more d a r k  autumn  i n v e s t i g a t o r s support  conchosporangial  are encountered  an  initiation  i s a major f a c t o r  Iwasaki,  winter Porphyra  of  is  of  response.  induction.  to  branch  with  of Dring  and  optimal  w o r k e r s on  induction.  results  no  conchosporangial branches  tested,  Richardson  photoperiod  1959;  that  under s h o r t d a y s .  conchosporangial nothing'  My  to those  in  nereocystis  working  response.  gardneri  a photoperiod  photoperiod  showed t h a t c o n c h o s p o r a n g i a l b r a n c h phytochrorae-mediated  under  s t u d i e s of most  indicate  (1967),  Porphyra  Sato  In  P.  (1967)  phase  umbilicalis. found  that  112  ccnchosporangial branches Iwasaki  and S a s a k i  subgrbiculata  were most a b u n d a n t a t  (1972),  Kjellman  i n an e x p e r i m e n t  forma  photoperiod rather  than the  (photosynthetic  effect)  16:8.  latifolia,  total  amount  which  with  showed of  was  Porphyra  that  light  i t was  received  responsible  for  photoperiod  for  ccnchosporangial branch formation. The  apparent  lack  ccnchosporangial  of  branch  an  induction  remains  an enigma.  formed  were  because  none o f them r e l e a s e d  premature  having  i s that  Other  Rentschler  be  The  no  induction  from  report  on  branches.  a higher temperature.  shown t h a t  a  i n the l i t e r a t u r e  of  conchosporangial  modifying  induction  did  appeared  i n those  formed  Hirano,  1956a, 1956b; I w a s a k i and  temperature  for  that  this  usually  over  (Kurogi  noted  changes d i d not the  conchospore  an o p t i m a l  (1956b) and  temperature  was  liberation.  a  cultures  not a l w a y s These  range  been of  temperature  Sasaki,  Kurogi  of  observed  I n many P o r p h y r a s p e c i e s i t has  Hirano  the  branches.  In Porphyra n e r e o c y s t i s I  t h e r e was  (1966)  on  u n t i l the appearance  but t h a t  K u r o g i and  linearis.  influence  temperatures,  Furthermore,  branch  effect  while temperature they  but  perhaps  conchosporangial  c o n c h o s p o r a n g i a l branches  and  is  of  branches,  branches  branches,  a  t h a t c c n c h o s p o r a n g i a l branches f i r s t at  the  have  photoperiodic  conchosporangial  that  ccnchospores t h i s only  that  conchosporangial  duration  out  nereocystis  e t a l . (1972) f o r P o r p h y r a  can  (1967) f o u n d  Porphyra  ccnchosporangial  effect  of Bird  factors  photoperiodic  induce  to  assumption.  formation  in  I t must be p o i n t e d  assumed  phctcperiod  optimal  and  1972). Akiyaaa  the o p t i m a l are  general  113  trends  and  species. P.  should  F o r example,  umbilicalis  egually  Light effect  K u r o g i and  10°C  as a t  has  of  responsible  also  cnly  intensity  under  has  1969a, a s  P. c u n e i f o r m i s  Miura,  1961)  (Setchell  produce  ccnchosporangial branch  the  conchosporangia  assumed  that  functioned contents  as  of  a a  P.  P.  2-4  conchospores.  or a  level,  was  however.  West  ( l e s s than  was  12:12).  the  Abe  conchospore (1966),  y e z o e n s i s , observed Migita  that  the  conchospores  in  release. cf  has  in  been  releasing  (Conway  working  generally  entire  and C o l e ,  1977);  Porphyra  tenera  with  1974)  branch  its  each conchosporangium  (1967b,  either  ccnchosporangial by  have  from  observed It  as  (Krishnamurthy.  phase d i r e c t l y  not  of  I t o , 1959,  Krishnamurthy)  the formation  thuretij.  cell  single  and  and  Knaggs  t h i s response  miniata  e t Hus)  conchosporangium  however, M i g i t a and  (Kurogi  intensity,  ( M i u r a and  the f o l i o s e  was P.  each  as  that  without conchospore  i s unfortunate that  Eorphyra g a r d n e r i  modifying  Release  and  It  light  inductive daylengths  sp.  to  a  nutrient  p o i n t e d out  Poiphy.ra  reported  have  initiation;  Miura  been  that  tranches  In Rhodochortcn, that  Porphyra t e n u i p e d a l i s ;  reported  production  and  tetrasporangium  Conchospore  Only  branch  light  investigated  6.  been shown to  demonstrated  ( p e r s o n a l communication)  (1967)  20°C.  1967)  for  f o r a l l Porphyra  conchosporangial  1955b; Conway, 1964b).  1966b,  combination  Sato  formed  ccnchosporangial  1955a,  (1966a,  at  intensify  on  be assumed t o be t r u e  conchocelis  as w e l l  Hirano,  not  has  produced  subsequently  114  studied  this  discussion).  per  observation  as s u g g e s t e d  my  four  each  I was  USEEh-Y-E . 3 S E ^ S ® £ i 3  Porphyra  1966;  than  one  purpurea.  This  month,  was  of  site  meiosis  of  the  at  proved  Conchospore which  to  be  release  by  correlated  i n the  VII).,  field  when  In Table VII study  a r e b a s e d on o n l y  but t h e g e n e r a l t r e n d (Hollister,  two  1966).  release  conditions  is  free-living  Abe,  7-10°C,  point  produced  whereas  (Table  Kurogi  unable  1968),  and  species  and  to demonstrate  responsible P.  for  thuretii,  points to  release  K u r o g i and  a l . 1967;  provides  division  typically  temperatures  i t  interesting  ( M i g i t a and  optimal  appeared  triggering  1963;  more  a  single  i s t h e same a s a  nearby  site  measurements a r e made.  temperature  factor  An  consistently.  was  first  Although  other  for  the c o n c h o s p o r a n g i a l branch  conchospore  f o r Amphitrite Point  where d a i l y  et  produced  s u r f a c e seawater  that  t o form  w e l l with temperature  measurement  the  conchosporangium  to t r i g g e r  gardneri  site  that  conchocelis  experiments  Porphyra g a r d n e r i  al.  division  per  conchocelis often  reasonably  IV  more  f o r Porphyra  by Conway e t a l . ( 1 S 7 5 ) .  conchospores  impossible  Part  illustrated  evidence  shell-living  In  (see  t o form c o n c h o s p o r e s i s t h e  not the f i r s t  that  (1961b)  conchosporangium  circumstantial  and  detail  i s of c o n s i d e r a b l e importance because  conchosporangium  £•  further  Kornmann  conchospore  good  in  1956a,  A k i y a m a , 1966;  B i r d e t a l . 1972;  conchospore work which  and  release  as  Chang,  1956b;  the 1956,  Migita  K u r o g i and S a t o ,  Shinmura,  that by  h a s been done  temperature  (Tseng  Hirano,  conclusively  1974;  on  primary Tseng and  et Abe,  1967;  Kurogi  Migita,  1974;  115  Chihara,  1975).  Migita  (1974)  without changing  the  conchospores  even  temperature.  My  release  may  reported medium  when  c o n c h o c e l i s phase  for  i t  failure  be  that 3  was  to  months  failed  transferred  obtain  attributable  to  cultured to  form  to  an i n d u c i n g  consistent  ccnchcspore  such  a  nutrient-related  phenomenon. In contrast observe  any  history  release. has  direct  phases  intensity  a  to  or  most  reports,  correlation  cf Porphyra continuous  Other  workers  modifying  have a l s o on  1955a, 1955b; S a i t o ,  Iwasaki  and  S a s a k i , 1972;  reported  morning  that  (Yamasaki,  1955b; T s e n g  and  Photoperiod release. Sato al.  The  (1962a, (1967)  Other ccnchospore desiccation  by P.  Kurogi  the  life light  conchospore intensity  ( K u r o g i and  akiyama,  1965;  K u r o g i e t a l . 1962).  Several  workers  liberation  e t a l . 1954;  i s h i g h e s t i n the  K u r o g i and  Hirano,  1956). secondary  factor  mediating  (1959), I w a s a k i  Kurogi  shown  photoperiod  liberation  a  1962b),  ccnchcspore release. that  1956;  release  not  and  work o f K u r o g i  has  ccnchospore  1954b; S u t o  is  inhibited  did  low  reported that l i g h t  ccnchospore  Chang,  and  He d i d n o t e t h a t  illumination  effect  (1961)  between t e m p e r a t u r e  tenera.  Hirano,  have  Iwasaki  et  (1961),  a l . (1962),  and  conchospore K u r o g i and Kurogi  ejt  t h a t t h e r e i s an o p t i m a l p h o t o p e r i o d f o r In c o n t r a s t .  variation  Bird  did  et  not  a l . (1972) affect  found  conchospore  linearis.  factors  t h a t have b e e n r e p o r t e d t o have an e f f e c t  release are a g i t a t i o n (Graves,  1969).  (Tseng and  Saito  Chang,  1956)  (1955) c o r r e l a t e d  cn and  vertical  116  turbulence in  i n the water column  the s u r f a c e  (1956a)  both  liberation reported  Kurogi  reported  a  (1953b)  1-2  my a t t e m p t s  nereocystis  Woessner,  recently  J u d g i n g from combination intensity  of  are  temperature  induce  of  tides  as  reports i n the specific  being of primary culture  studies  have a t h o r o u g h branch  be  the  best  i t responded  photoperiod  to  faster  1977)  have  by s u b j e c t i n g  I  think  that  the  conchospore  the  and  light  release,  with  much  more  critically  a b l e t o m a i n t a i n must be  done  understanding of the process of  From my  one  California  photoperiod  under  induction  the 3 s p e c i e s I s t u d i e d .  by  importance.  will  conchosporangial  release  literature  for  before  we  in  August,  temperature,  important  „•  release  12°C.  c o n d i t i o n s t h a n I was  of  Hirano  i n conchospore  conchospore  workers  triggering  controlled  Of  and  i t with s p r i n g  communication  in  a l l  Further  7.  conchospores  Kurogi  week p e r i o d i c i t y  failed,  personal  succeeded  to  c o n c h o c e l i s t o a temperature  because  of  by T a k e u c h i e t a l . ( 1 9 5 4 ) .  Porphyra  would  and  but were u n a b l e t o c o r r e l a t e  although  (J.  water.  w i t h an i n c r e a s e  and  release  experience, Porphyra  subject  and  conchospore  more  to further definitely  regimes than d i d t h e . o t h e r 2  in  thuretii  investigation to  different  species.  Conchospores  particular  amoeboid change  jgrphyra t h u r e t i i . Bhodcphyta,  interest c f shape These  was and  two  having p r e v i o u s l y  the photographic movement  by  conchospores  phenomenon a r e n o t been r e p o r t e d  documentation of  unknown i n the  f o r : monospores o f  117  Banjgia  (Beinke, c i t e d  from  1882;  Kylin,  Sommerfeld  (Berthold,  1922;  1882; K u n i e d a ,  lOlilociadia (Svedelius,  1945,  and  and  Ljagpra  p. 431;  Nichols,  1939; K u r o g i ,  (Nichols  1917),  (Rcsenvinge, Borsje,  Fritsch,  (von S t o s c h ,  Stegenga  and  Graves,  1967),  Vroman,  1976);  for  (Conway a n d  for  1939);  SQI.2h.2LSi Graves,  1969),  Bangia  (Thuret  Fritsch,  Bonnemaispnia (von  (Kunieda,  Stosch  (Chemin, 1965),  p..,.602),  and  1929); and  Scinaia  Kylin,  Nichols,  1922; 1970),  (Kuckuck,  (Chemin,  cited  1927),  and f o r t e t r a s p o r e s  Acrochaetium  1973);  f o r carpospores of  and B o r n e t , 1 8 6 7 ) , Platoma  1945,  #  (Sommerfeld  released  Cole,  ( J a n c z e w s k i , 1873; Okamura e t a l . 1920;  Helminthora from  spermatia  Acjcochaetiuro  1929; S t e g e n g a and  • c a r p o g o n i a ' o f P o r p h y r a pa p e n t u s s i i Porphyra  1969),  Helminthora  1965) and  1909, a s C h a n t r a n s i a ; D a n g e a r d ,  1976;  1970), P o r p h y r a  1961;  Lissant,  Berthold,  of  and  Liagora  ( S t e g e n g a and Vroman,  1976} . Some r h o d o p h y c e a n amoeboid  changes  reported  by  spores exhibit  of spcre  Dangeard  shape.  (cited  gliding  Such  Frythrotrichia,  Dumontia, spermatia 1945,  changes  also  been  for  Recently  Porphyridium  slight has  Phyllophora;  p. 602)  Ggniotrichum. °f  tetraspores  and  of  L i n e t a l . (1975)  o f shape. observed  exhibited They in  been  1945, p. 602)  (1927)  and by Chemin  monospores  which  has  for  for  monospores  o f A n t i t h a m n i o n . C e r a m i u j , and  and c a r p o s p o r e s o f Ceramium. of  locomotion  from F r i t s c h ,  A c r o c h a e t i u m c a r p o s p o r e s ; by B o s e n v i n g e °f  movements w i t h no  (cited  from  and  Fritsch,  Erythrotrichia made a d e t a i l e d  gliding  reported  Polysiphonia,  and study  movements w i t h o n l y  that  this  Chrpothece m o b i l i s  phenomenon Paseher  and  118  Petrova  and i n o t h e r  Pcrphvridium  Amoeboid c h a n g e s o f s h a p e have  previously  1972)  P. a n g u s t a . P. k u n i e d a i  et and  Bangia The  out  that  this  (Scmmerfeld purpose  a  between t h e s e Neither movements light  had an  t h e B a n g i o p h y c i d a e and or  i s there  function  nor  known, a l t h o u g h B e r t h o l d A  polar  it the  probability  conchospores  may be t h e c a s e .  and  The  Porphyrjdium  expulsion  Is  some e v o l u t i o n a r y  link  of  the  published  spore  amoeboid  The  was  to  a suitable  (1972) of  was a  t h e moving c e l l s ,  that t h i s  be  photcgraphs  study  there  out that  could  finding  by M i g i t a  showed t h a t  established  the  function  ultrastructural  from  of  (1882) p o i n t e d  possible  of mucilage  was n o t d e f i n i t e l y movement.  Nemaliales.  mechanism  f o r attachment and g e r m i n a t i o n .  on  1972),  two g r o u p s o f H h o d o p h y t a ?  substrate  (1975)  (McDonald,  (Chen  a l l o f t h e s e examples i s t o p o i n t  the  al.  (Migita,  1970) .  increase  this  conchospores  1961), P. m i n i a t a  e t Abbott  and N i c h o l s ,  influence.  P. y e z o e n s i s  by  f o r Porphyra vezoensis  Hollenberg  coincidence  the  is  movement  (Kurogi,  of l i s t i n g  most a r e from  just  and  been r e p o r t e d  a l . 1970), P. s m i t h i i  species.  the  cf  suggest Lin et  production although cause  of  119  Table VII. A n n u a l t r e n d i n s u r f a c e w a t e r t e m p e r a t u r e (°C) a t t h e s t u d y s i t e (48° 50.1 !*, 125° 11.1*W) and A m p h i t r i t e P o i n t (48° 55.2*N, 1 2 5 ° 3 2 . 2 * W ) . 1  Study January February March April May June July August September October November December  site  7.5 7.0 7.0 10.0 12.0 12.5 12.8 13.8 13.5 11.3 9.0 8.0  Amphitri  ——— 8.5 8. 1 9.6 10.0 11.7 12.3 12.2 12.2 11.2 9.9 9.2  120  Porphyra Figure Figures  12a,  gardneri  Monospores being r e l e a s e d along the margin of a t h a l l u s c o l l e c t e d a t the study s i t e .  12b-f. Various stages i n the development c f the f o l i o s e t h a l l u s from a monospore. Note the lon< r h i z o i d a l protuberances.  Figure  12g.  Released  carpospores.  Figure  12h.  Carpospore germination phase.  Figure  12i.  S i n g l e s p o r e - l i k e s w e l l i n g on the filament.  Figure  12jj.  Conchosporangial branches.  to g i v e the  conchocelis conchocelis  121  122  Porphyra  gardneri  Figure  13a.  Conchocelis filaments, ccnchosporangial (lower r i g h t ) and r e l e a s e d c o n c h o s p o r e  Figure  13b.  Bipolar sporeling resulting germination.  Figure  13c.  C h a r a c t e r i s t i c p i n n a t e l y b r a n c h e d morphology o f t h e c o n c h o c e l i s phase g r o w i n g i n o y s t e r s h e l l . Porphyra  from  branch (arrow).  conchospore  nereocystis  Figure  13d.  Released  carpospores.  Figure  13e.  Carpospore germination phase.  Figure  13f.  Basal s p o r e - l i k e s w e l l i n g of the phase.  Figure  13g.  A p i c a l s p o r e - l i k e swelling of phase.  Figure  13h.  I r r e g u l a r and  beaded-branch  Figure  13i.  Sguare-celled  branch.  Figure  13j.  Small ccnchosporangial v e g e t a t i v e f i l a m e n t by  Figure  13k.  Ccnchosporangial  to give  the  conchocelis  conchocelis  the  conchocelis  types.  branch connected t o the a s q u a r e - c e l l e d branch.  branches.  123  124  Porphyra n e r e o c y s t i s Figure  14a.  C l o s e u p of c o n c h o s p o r a n g i a l branches s i n g l e s t e l l a t e c h l o r o p l a s t i n each conchosporangium.  Figure  14b,  P i t p l u g s (arrows) between c c n c h o s p o r a n g i a i n t h e c o n c h o s p o r a n g i a l b r a n c h ( t r e a t e d w i t h chromosome f i x a t i v e and s t a i n ) .  Figure  14c.  Divisions  Figure  14d.  C h a r a c t e r i s t i c pinnate branching of the c o n c h o c e l i s phase g r o w i n g i n o y s t e r s h e l l . the i r r e g u l a r s w e l l i n g s .  within the conchosporangial  Porphyra  showing  the  branch. Note  thuretii  Figure  14e.  Released  carpospores.  Figure  14f.  Carpospore phase.  Figure  14g.  V e g e t a t i v e c o n c h o c e l i s f i l a m e n t s , beaded b r a n c h e s and young c c n c h o s p o r a n g i a l b r a n c h ( a r r o w ) .  Figure  14h.  Cluster of conchosporangial tranches s i n g l e s t e l l a t e c h l o r o p l a s t i n each conchosporangium.  Figure  14i.  C a n d e l a b r a - s h a p e d c l u s t e r of c o n c h o s p o r a n g i a l branches showing t h e i r p o s i t i v e phototropism.  Figure  14j.  C c n c h o s p o r a n g i a l branch  Figure  14k.  Released  germination  conchospores.  to g i v e the c o n c h o c e l i s  with  showing  conchospores.  125  126  Porphyra Figure  Figure  Figures Figure Figures  15a-d.  15e.  thuretii  C o n c h o s p o r e s e x h i b i t i n g amoeboid change o f s h a p e and movement. F i g u r e s 15a and 15b t a k e n seconds apart. F i g u r e s 15c and 15d t a k e n 45 seconds apart., Young b i p o l a r s p o r e l i n g r e s u l t i n g from conchospore germination. Note r h i z o i d a l protuberance.  15f-15g. Older, uniseriate sporelings starting form b a s a l pad. 15h.  90  B i s e r i a t e stage  15i-15j. pad  to  of s p o r e l i n g growth..  Older s p o r e l i n g s . i n F i g u r e 15j.  Note t h e b a s a l  rhizoidal  127  128  PART IV  - CYTOLOGICAL STUDIES  General The  Introduction  first  three  of  the  morphological  Porphyra g a r d n e r i ,  P.  n e r e o c y s t i s and  purpose  IV  discussion  of  histories  Part  of these  w i t h e m p h a s i s on reproduction I  is  t h i s t h e s i s presented  to  3 species  in i t s life  and  i n as  to  P.  nereocystis  u l t r a s t r u c t e r e of  Cytological studies site.  The  were  £•  and  thuretii  chromosome  ethanol;glacial  counts  occurring  count and  life  possible, sexual  could  because  because the  attachment  to t h e i r  host  be of  of  plants,  i n P.  material  in  and  gardneri.  c o l l e c t e d at  gardneri May,  used  and  the  in  the  those  of  i n January. was.  material (3:1),  aceto-iron-haematoxylin-chloral  chrcmoscffle  primary  Methods  collected  acetic acid  Spermatangial material  as  occurrence of  the  of Porpfryra  study  For  And  were done on  specimens  P.  detail  the  The  cf  cytological  monosporogenesis  fertilization nereocystis  the  much and  histories  thurgtji.  examine  Materials  study  P.  a detailed  history.  wanted  £orphyra g a r d n e r i  life  document  Porphyra gardneyi  also  observe the  parts of  and  hydrate fixed the  a t any large  chromosomes i n the  fixed  in  stained (Wittmann, time to  95% with  1965). obtain  a  number o f d i v i s i o n s mature  spermatia  129  are  condensed.  The  best  chromosomes were b e g i n n i n g chromosomes  in  the  because they  tended  counts  were made on  type  was  the  determine at  (1S70)  to  and  at  (Kito,  Cultured hours.  Dividing cells to the  material fixed  Sommerfeld  phase  also  highest  i n the  In  an  metaphase, 0.1%  and  arrest  and  the  and  at  give  hour  for  T h i s was  another  mitosis  and  Tokida,  and  of  1967;  most  Migita,  of  the  i n Part I I I .  intervals  for  conchocelis  24  phase  were f o u n d first  in few  Pseudogloiophloea Bangiaconchocelis cell  division  was  period.  to  conchocelis  Austin  1971).  growth  Most d i v i s i o n s  of  begin  occurs  Mclachlan,  freguency  to  to  Porphyraspecies.  that  hourly  hours  found  Pringle  (1970) i n work on  dark  material i t  24  p e r i o d , p r i m a r i l y i n the  the  reliable  gardneri.  Nichols that  condensed  of P o r p h y r a  were i n f r e g u e n t i n t h e  dark  the  chromosome  conditions  fixed  as  Spermatial  (1969) i n work w i t h  attempt  increase  the  number  of  cells  phase c u l t u r e s were t r e a t e d w i t h  1$ c o l c h i c i n e f o r 24  metaphase i n h i g h e r  however, i t had  not  Ramus  found  during  yabu  foliose thallus.  hours of darkness. and  did  i n c u l t u r e have been d e s c r i b e d  c o n c h o c e l i s was  compared  highly  K i t o , Ogata and  Methods o f i s o l a t i o n phase  with  emphasized  night  just  The  hours.  results  1967b; Yabu, 1969a, 1970;  conchocelis  every  1-2  obtained  carposporangial  occurring.  for  have  plants  and  thalli  similar  workers  freguently  clump.  locality  fix  continue  obtained  Several  to  when m i t o s i s was  sunset  spermatia  case of v e g e t a t i v e  necessary  were  to condense.  mature  counts  In  counts  no e f f e c t  hours.  plants on  the  T h i s compound (Klein  conchocelis  and  0.051?,  i s known t o  Klein,  phase.  in  1970);  130  Material fixed For  and  attachment  embedded  Eeulgen  thick  for the  in  staining,  were  cut.  solution  (Dimedone)  for  then r i n s e d  i n running  1968).  18 h o u r s  Slides  potassium  a t rocm t e m p e r a t u r e .  carried  running  followed  water  for  10  counterstained  for  1  Green  ( F e d e r and  in  phosphate  dehydration Sections  ethanol  minutes  run  pH  on  1:3),  0.5  7.2  Specimens coated  and  in  were  (1:1)  O'Brien,  60°C  Sharma,  for  20  reagent  1965)  washes  for 3  of  were  0.5% in  lightly  with  1 hour, with  using  citrate  electron  point a  dried, Cambridge  Om  03  in  50$  1963)  and  microscope.  Thallus  in  resin.  fleichert  (Reynolds,  of ethanol:amyl hr  OsOy  f o l l o w e d by  Spurr's a  microscopy  procedure.  0.5  i n 2%  with uranyl acetate  lead  and  critical  postfixed  for  knives  EM-10  series  each  examined  at  were  i n each), then a r i n s e  infiltration  and  a Carl Zeiss  hr  slides  to S c h i f f " s  and  m a t e r i a l was  glass  1971)  a graded  using a  1968) .  u s i n g t h e above  through  The  Sections  Material f o r scanning electron dehydrated  HCl  successive  T h e y were s t a i n e d  (Dawes,  examined w i t h  3  minutes.  i n e t h a n o l and  ultramicrotome.  of Sharaa  by  microscopy  cut  um  m i n u t e i n a 1% aqueous s o l u t i o n o f F a s t  buffer  were  was  3-6  out  ( F e d e r and  transferred  (2  O'Brien,  For e l e c t r o n  out i n 1N  then  metabisulfite  1977)  carried  water f o r 30 m i n u t e s  were  T h i s was  staining  5,5-dimethylcyclohexane-1,3-dione  (made a c c o r d i n g t o t h e r e c i p e hours.  (Henry,  b l o c k a d e was  of  H y d r o l y s i s was  minutes.  sections  aldehyde  saturated  Feulgen  m e t h a c r y l a t e as d e s c r i b e d i n P a r t I I .  serial  fin  s t u d y and  was  fixed  and  p i e c e s were  then  acetate  100%  (3:1,  amyl  mounted on  1:1,  acetate. stubs,  Stereoscan  Type  gold 2k  131  scanning  a.  electron  Epiphyte-'Host  microscope,  Attachment  1  Introduction The summarized  known  epiphytic  by Ohmi  (1963) and T c k i d a  epiphyte-•host* received  relationship  little  attachment  of  laciniata)  to  penetrated rhizoids deeply  Porphyra Fucus  the  host  these  reported  that  Porphyra  tissue  var. porracea  those  rhizoids  the  basal  sp.) p e n e t r a t e d  of  i t s  (Mertens)  of  has  P. u m b i l i c a l i s v a r .  (1966) (as  of the  described the  the basal  Hiura  been  species  that  Kito  whereas  (1923)  (as  have  The n a t u r e  Porphyra  Grubb  that the basal r h i z o i d s  tissue  Porphyra  concluded  the  Grateloupia filicj,na  of  (1960) .  purpurea  and  P^ k a t a d a i  into  (1968) found  in  investigation.  host c e l l s . of  species  o f P. o n o i  host  Howe.  Fukuhara  Ueda  penetrated  P. p s e u d o c r a s s a  Yamada  et.  Hikami d i d not. There  have  epiphyte-* host The host  that As  restriction  they the  into may most  relationship study  no  interaction  1  s p e c i e s and  develop  been  physiological  studies  i n the epiphytic Poiphyra  of  species.  of Porph vra n e r e o c v s t i s t o e s s e n t i a l l y  the  failure  mature t h a l l i  of  i n culture  have a p h y s i o l o g i c a l probable  P. g a r d n e r i  region  dependency  made.  III)  on  the  one to  suggests »host . t  o f n u t r i e n t exchange i n such a  would be t h e a t t a c h m e n t  o f t h i s r e g i o n was  sporelings  (see P a r t  the  zone, a  light  microscope  132  Results  1.  Porphyra  gardneri  Transverse  s e c t i o n s were c u t t h r o u g h t h e b a s a l  £2ifihyra g a r d n e r i rhizoidal £•  and  filaments  gardneri  spread  the  central portion of  a b a s a l pad {Figure  medulla  (Figures  penetrate  to v e r i f y  2»  Porphyra  Sections  surface  filaments  extensive  cells,  cut  The r h i z o i d a l  penetrate  margin  filaments i n  deeply  into  the  appear t o  a l t h o u g h an E. N. e x a m i n a t i o n i s  through  spread  16c)., I  n  stipe  rhizoidal  the  basal  and t h e s t i p e o f N e r e o c y s t i s  out to  This the  make a s l i g h t  Nereocystis  blade  nereocystis  3§£^II€£i«  filaments  pad  The  c f the t h a l l u s of  on t h e L a m i n a r i a  to the s t i p e a x i s .  (Figure  base  margin.  this.  Porphyra n e r e o c y s t i s  rhizoidal  blade  16a and 1 6 b ) , and i n some c a s e s t h e y  were  perpendicular  the  16a).  this  the medullary  needed  £•  form  out s l i g h t l y  form  y  Laminaria  which  to  fa  the  portion of  region in  a  plane  &s i n P. g a r d n e r i . t h e b a s a l form  a  pad  on  the  pad i s much l a r g e r t h a n t h a t center  indentation  (Figures  16c  penetration  of  of  the  into and  pad the  16d),  stipe formed  the rhizoidal cortex  but  of the s t i p e t i s s u e .  of  there  the  i s no  133  Discussion  The  intimate  Laminaria  blade  attachment suggests  metabolites  from the  by  et  liining  In E«  Laminaria  contrast  to  penetration  exchange  of  of  material.  compounds £• of  nergpcystis. P.  nereocystis  development and  may  the The  such a s  the  (1973b)  a l s o be  water  mechanism may  be  lack  of  Studies Laminaria  attachment of  of  significant  there  zone.  may  not  be  I t should  be  that  a  p r e r e q u i s i t e f o r exchange Nereocystis  they  are  d e t e r m i n i n g the  acting  transfer  demonstrated  that  where  the  tissue.  the  attachment  a  a  most  medulla  the  t i s s u e i s not  case  into  at  Harlin  host  The  that  gardneri. and  be  of s t i p e t i s s u e s u g g e s t s  that  of  may  to  Porphyra t h a l l i .  shown  i n the  Porphyra  materials  though,  penetration  to the  is superficial,  rhizoidal  Porjahjrra g a r d n e r i there  have  are t r a n s p o r t e d  ngreocystis  noted  that  a l . (1972)  photosynthates  of  at  an  is  picked host  time o f i n i t i a l  and  physiological studies  up  by  specificity  earlier  the  exuding  stage  conchospore  in  settlement  germination. Oltrastructural  elucidate  the  and  i n these o b l i g a t e  "host*  nature  of  the  are  needed  a s s o c i a t i o n between t h e  e p i p h y t i c Porphyra  to  epiphyte  species.  134  B.  Monosporogenesis  Introduction S r o i t h o r a naiadurn Hosenvinge  are  the  has  not  and  been a n y  Por.phyra, the  Cole,  1971;  cycle  fact  (Areschoug)  level  respectively).  There  investigation  commercially decided  to  observations  o f monospores i n importance  cultivated make  of  i n which  ultrastructural  i n view o f t h e g r e a t  in  therefore  ultrastructural  c^ccinea  McDonald, 1972,  ultrastructural  monospore  Porpfeyra  been e x a m i n e d a t an  a surprising  s p e c i e s . , I t was  Porphvropsis  o n l y members o f t h e B a n g i o p h y c i d a e  mcncsporogenesis has (HeBride  and  some  of  Porphyra  preliminary  monosporogenesis  in  gardneri.  Results Monosporic t h a l l i their  gelatinous  Under t h e larger had  light  than  fine  margins  vegetative granular  the  cells  and  contents  (see  The thalli  than  the  u l t r a s t r u c t u r e of was  examined  by  (see P a r t I ) . were  slightly  the s t e l l a t e Tanaka  They  chloroplast  (1952)  o k a m u r a i Ueda had  also more  was  noted  granular  cells.  the in  monospores  field  more s p h e r i c a l i n shape.  Part I I I ) .  vegetative  i n the  monosporangia  and  t h e monospores o f P o r p h y r a  contents  recognized  releasing  microscope  somewhat o b s c u r e d that  were e a s i l y  distal  p o r t i o n of  transverse  monosporangial  section.  The  first  135  indication  that  transition  to  activity of  the  monosporangia  (Figures  small  vegetative  17  and  fibrous  association  with  monosporangium  was  19a)  a  central  starch  g r a n u l e s were a b u n d a n t . prolifically  releasing  margin)  to  be  monosporangia vesicles (Figures the  appeared 18  and  light  19c)  as  In large  material the  to  as  inner  cell  wall,  chloroplast  with  Floridean vesicles  (those toward fibrous  The  the  vesicles in  small  the  fibrous vesicles  their contents  between  resulting in a layer  of  19b).  v e s i c l e s increased  with  Each  fibrous  (Figure  monosporangia  that  were  number o f  19c),  v e s i c l e s became e x t r a c y t o p l a s m i c .  associated  in  19a).  fibrous  form l a r g e r  release  found  observed  more mature m o n o s p o r a n g i a t h e  fibrous  fibrous  granules  microscope l e v e l .  well  (Figure  number o f  T h e s e abundant  to coalesce  plasmamembrane and  fibrous  18).  production  nucleus.  monosporangia  numerous  at the  stellate  a  dictyosome  were  (Figure  lateral The  in cider  (Figure  the  to  in  i n the  Mitochondria  single  pyrenoid  undergoing  increase  dictyosomes  contained  increased  were  which r e s u l t e d  central  appear  an  vesicles. these  and  cells  and  This  small the  and large  phenomenon  near  the  was  releasing  margin. Unfortunately were  observed,  dissolution  of  the  only and cell  the  initial  were wall  s t a g e s of  monospore  characterized layers  beneath the  by  an  outer  release apparent  cuticle.  136  Discussion Monosporogenesis the  i n Porphyra g a r d n e r i i s c h a r a c t e r i z e d  production of small  to  be  produced  ultrastructura1  the  as  Dltrastructurally reported  1971)  and  in  in  tetraspores  (see McBride  different  types In  picture  Part  no  and  mucilage  layer  1970)  and  appears  chemical  nature  (1971) r e p o r t e d  vesicles  during  lack  Another  has  to  be  a  aid  the  for  vesicles.  a  in  Such  complete  t h e one  in type,  monosporogenesis. wall  (observed,  by a l a y e r  i n the f i b r o u s  needed produced  monospore  but  of mucilaginous vesicles. (Peyriere,  adhesion to  to  the  determine  the  by  the  needs t o be examined of  and  the p r o d u c t i o n of 2  monospore is  as  Cole,  carpospores  mucopolysaccharide  material  a s p e c t which  t h e same  monosporogenesis  cell  i t s origin  o f enzymatic d i g e s t i o n .  fibrous  a  are surrounded  w a l l breakdown a t t h e t i m e  result the  Cole  may  of  (1970) f o r  ( M c B r i d e and  spermatogenesis,  Histochemical study  vesicles. cell  IV,  doubt  This  dissociated  Dixon  spermatia,  (1972) i n P o r p h y r p p s i s  omitted)  substrate.  monosporangia  algal  monospores  m a t e r i a l which  have  Porphyra g a r d n e r i I observed only  as d i d McDonald Released  which  Porphyra.  red  and  This  monospores a r e more  has b e e n s u g g e s t e d by  Smithora  of  that  appear  activity.  t h e f i b r o u s v e s i c l e s appear  other  summary).  Smithora.  dictyosome  vegetative c e l l s  B a n g i a and  those  by  s t u d y has d e m o n s t r a t e d  thallus,  monospores o f  l a r g e f i b r o u s v e s i c l e s which  primarily  than u n d i f f e r e n t i a t e d frcm  and  by  i s whether  release  enzymes may  be  fibrous  is  released  the by  137  C.  Sexual  Reproduction  In Porphyra  gardneri *  Introduction  The  occurrence  Porphyra  has  reported  by  several  Kunieda,  Berthold  1921; 1939;  1972;  Tseng  Yabu, no  workers thought  1969;  Conway  e t a l . 1975).  types  of  referred terms  that  al.  Dixon,  1970,  spore  and  Hawkes,  and  1967b;  and  Kurogi, and  (1902) r e p o r t e d  reproduction,  Conway,  Conway and  1927;  Giraud  f o r i t was  and  inconclusive  1964a,  Cole,  (1964a) s u g g e s t e d  1964b;  1973,  1977;  t h a t because of  of s e x u a l r e p r o d u c t i o n the  carpospores.  to t h i s ,  and  t h e terms  O^-spore  1977b)  first  Chang, 1955;  sexual  t o r e p l a c e spermatium  1973)  I t was  Dangeard,  formed i n •packets' i n Porphyra  t o as s p e r m a t i a  (Dixon,  and  1959 ;  1973;  Conway  1924;  evidence  of c o n c l u s i v e evidence  -spore  1975;  the  genus  leucosticta,  However, Hus of  the  other species concurred  Migita,  196 9b).  evidence  carpospore. , In a d d i t i o n cell  1963;  Krishnamurthy,  Graves,  lack  of  1952;  could find  the  Porphyra  Hagne,  and  1954a;  for  investigators  Yabu and T o k i d a ,  in  phycologists.  Hamel,  that  (Drew,  (1882)  by  1924;  1968;  other  reproduction  Grubb,  Magne, he  sexual  long been debated  subseguent  (Ishikawa,  1961,  of  have  mother been  She  s h o u l d not proposed  c^-spore  used  *This portion o f t h e t h e s i s has been a c c e p t e d i n P h y c o l o g i a and i s ' i n p r e s s * i n V. 17(3) .  for  be the  to replace  'carpospore* cell  two  mother  (Conway  et-  instead  of  publication  138  carpogonium .  The  1  (McDonald, to  has  met  with  1972) b e c a u s e t h e n o n - p i g m e n t e d c e l l s  criticism  that i t refers  do n o t g e r m i n a t e and a r e t h e r e f o r e n o t s p o r e s . The  of  J& - s p o r e  term  majority o f c y t o l o g i c a l  sexual  reproduction.  chrcmoscme  number  for  subseguent  workers  carpospores  in  chromosome cells  counts  Magne the  have  22  s t u d i e s have p r o v i d e d  other have  (1952)  reported  carpospores  obtained  of  (Table  a  diploid  P. l i n e a r i s  diploid  species  evidence  counts  and  f o r the  VIII).  Diploid  a l s o been r e p o r t e d f o r t h e v e g e t a t i v e  o f t h e c o n c h o c e l i s phase i n  9  Porphyra  species  (Table  VIII) . There meiosis. Tseng  i s seme d i f f e r e n c e o f o p i n i o n r e g a r d i n g According  and  Chang  carpogonium however,  i s  the  which  occurrence  and  purpurea  the  Giraud  (as  of (1959)  Dangeard  division  Magne  are  haploid,  (1968) and K i t o  of meiosis.  have been p o i n t e d sexual  t o as e v i d e n c e  reproduction  report that a l l l i f e  P. u m b i l i c a l i s  a r e p o r t by Conway and C o l e  and  of the f e r t i l i z e d  carpospores and  (1927)  t c form t h e conchospores i n the  branch i s the s i t e  Krishnamurthy^ !•  and  that the division  data  (1921),  the f i r s t  (1967b),  ccnchosporangial The  (1955) meiotic  Migita  (1974) f o u n d  t o Ishikawa  the s i t e of  var.  consists history  laciniata)  (1973)  that  against of  stages of  are haploid*,  the  conchocelis  T h e s t u d i e s which a r e being r e p o r t e d i n t h i s thesis indicate that there i s no l o n g e r any r e a s o n t o c o n t i n u e t o u s e t h e t e r m i n o l o g y p r o p o s e d by Conway (1964a) f o r Porphyra g a r d n e r i . The terms spermatium, carpogonium, prototrichogyne and c a r p o s p o r e s , w h i c h have p r e v i o u s l y been applied t o Por P h y r a . will be used. See t h e t e r m i n o l o g y section f o r further comments. l  I n c o n t r a s t K i t o e t a l . (1971) o b t a i n e d the c a r p o s p o r e s o f t h i s s p e c i e s .  2  a  diploid  count  for  139  phase  o f P. p a p e n f u s s i i i s h a p l o i d .  (1873) t h a t the to  both c a r p o s p o r e s  same c a r p o g o n i u m express  1.  doubt  and s p e r m a t i a c a n  l e d Drew  that  they  The r e p o r t s o f J a n c z e w s k i  (1954a) and Conway e t a l . (1975)  play a r o l e  i n sexual reproduction.  step i n attempting t o c l a r i f y  nature c f t h e spermatium  in  considerable importance.  Krishnamurthy  similarity  of  mature  Florideophycidae,  and  Spermatangial  pointed  and  Bornet,  Tanaka,  cf  contradictory  that  the  thallus) that  (1921)  (1S56)  division  pointed  the  o f each  be  for  been  studied  of  further  by s e v e r a l  (Janczewski,  1959, 1972).  1873;  These  (1873),  and  first  (in the  to  (1902) r e p o r t s  give  four  spermatangium  are  division  plane o f the (1959)  that  spermatangia,  various interpretations  thought  there  is  a  and t h a t t h e  was p e r i c l i n a l .  to the d i f f i c u l t y i n d i s t i n g u i s h i n g  data  T h u r e t and B o r n e t  t h a t the  periclinal  the  (1952) and K r i s h n a m u r t h y Hus  studies  t h e s p e r m a t i a a r e fornsed by  maintained  o u t , these  vegetative c e l l .  to  with those i n the  need  spermatangium,  was  whereas T a n a k a  division  seem  (1959) commented cn t h e  microscope  Janczewski  i t was a n t i c l i n a l .  first  a  the  spermatangium  •cruciate^  due  light  has  indicate that  repeated d i v i s i o n  of  out  problem, the  1878; B e r t h o l d , 1882; Hus, 1902; I s h i k a w a ,  l i t t l e more t h a n  (1878) and I s h i k a w a  would  o f Porp.hy.ra  1952; K r i s h n a m u r t h y ,  in  this  o f t h e spermatium.  development  investigators using the Thuret  Porphyra  spermatia  cytological investigation  did  from  The Spermatium  as a f i r s t  1921;  originate  as  Drew  a r e perhaps  t h e spermatangium  from  140  Little  i s  known  about  spermatiufii i n P o r p h y r a . except  f o r the  carpospores. and  formed  by  and  little  repeated  flagne  tend  spermatium  mentions t h a t  is  in  surprisingly  numerous spermatium  1972  -  (Brown,  £tilota;  Polyneura  indicated  that  formation  and  vesicles,  and a  complexity  the  1977  made  P e y r i e r e (1974) abundant  general  Scott  1971,  to this,  genera  of  Kugrens,  and  1974 -  P cjLj sipjb_Q n i a,  1972a,  1973b -  1972b These  of  production chloroplast  which have c h l o r o p l a s t s ) .  of the  Kugrens  and Dixon,  features  in  1972  made  Bonnemaisonia).  the  studies of  been  Batrachospermum:  reduction  fibrous  contrast  now  several  are  of the  (McBride,  In  have  characteristic  (1977b)  nucleus  and E r y t h r o c v s t i s :  -  they  leucosticta.  and F u r c e l l a r i a :  maturation  ( i n those  the  1971, 1974 - G r i f f 1 t h s i a,  Young,  as  ( 1 9 5 9 ) , Hawkes  Bangia).  1969 -  the f a c t  pigmented  seen  been  Simon-Bichard-Breaud,  Eonnemaisonia:  mention  state.  have  in  the  that they a r e  that  studies  Levrinqiella  Janczewskia j P e y r i e r e , laurencia,  -  development  Florideophycidae West*  have  from  on t h e f a c t  a condensed  1972  ultrastructural  them  o n l y a few u l t r a s t r u c t u r a l  spermatia  McDonald,  produce the  the spermatia  less  of Porphyra  that,  that  Krishnamurthy  G, and J . Feldmann  Bangiophycidae Smithora;  become  c f the  opinion  that  Most w o r k e r s  (1977) r e p o r t  v e s i c l e s i n the spermatia Sather  (1959) s u g g e s t e d  to  (1952),  the  differentiate  divisions.  Mumford a n d C o l e  mature  to  structure  of divisions  were homologous b a s e d  that the spermatia develop.  number  Krishnamurthy  carpospores  internal  Hus (1902) was o f  larger  spermatia, there i s  the  studies  spermatium of  fibrous  size  I n most  and cases  141  the  nucleus of the  Starch  is  nature  usually  spermatium  absent  in  is in a  the  condensed  state.  mature s p e r m a t i u m  {Young,  1977). I f t h e spermatium male  gamete,  as has  i n Porphyra  i s in fact  been c l a s s i c a l l y  1882),  siiilar  t o the s p e r m a t i u m i n t h e F l o r i d e o p h y c i d a e .  the  first  step  reproduction  2.  conclusively  1956) were  were  of  gametes  released.  Drew  on  thallus  which  neither  in  a  following  different presence or  A study  undertaken  determine of t h i s  information  i f  of as  sexual  alga.  and  that  until  has  my  o c c u r r e n c e of a s e x u a l  fusion.  (according  absence  o b t a i n e d no  into  that a  fusion.  a  had  been  evidence of  dealing  She a  that  with  noted  cell  the  Drew n o t e d  cf f e r t i l i z a t i o n of  Drew,  carpospores  they  in  carpogonium  and  to  such  workers.  Bangiophycidae.  this  after  the l i t e r a t u r e  spermatium,  mechanisms  fusion  (1959)  indicated  been with  have s u b s e q u e n t  the  not  work  Koschtsug  underwent  transformed by  ultrastructurally  t h e o p i n i o n t h a t t h e s p e r m a t i a and  Porp.hyra  is  fertilized produced  (1856)  (1956) summarized  reproduction reports  piece of  Krishnamurthy  f u s i o n s , and  a  Fertilization  demonstrated,  Solier  to  history  gardneri, i s the actual  D e r b e s and  be  g a r d n e r i was  attempting  o c c u r s i n the l i f e  most v i t a l  Porphyra  i n Porphyra  in  B e p o r t s Of  The  perhaps  (Berthold,  would e x p e c t  development  would  as  t h e n one  spermatium  i t  stated  functioning  sexual  that the  which  foliose is  then  carpospores  that  t h e r e were  d e s c r i b e d , based  prototrichogyne  most  produced  on by  are two the the  142  carpogonium.  She  prototrichogynes and  Kunieda,  without  1939  the  carpogonia  are  that  Bosenvinge  also  and  carpogonium,  i s  a  out  protruding),  Similarly,  fertilization  distromatic  canals  into  he s t u d i e d .  species,  attached  next  for  to the thallus  1896; B o s e n v i n g e ,  to  (1973)  spermatia  of  the which  r e p o r t e d , t h e r e i s o n l y one, l o c a t e d  (Joffe,  reported  spermatia  t o p o f them.  surface.  1909; D a n g e a r d ,  Kurogi,  carpogonia  1961)  canals.  via fertilization  suspected  Similarly,  laciniata)  spermatia i n  Krishnamurthy  s u r f a c e o f P. but  did  He d i d r e p o r t s e e i n g c a n a l s He s a i d  have  o f P. p a p e n f u s s i i b u t t h e y  on the t h a l l u s  var.  sheath  Dacgeard  pole  fertilization  mucilaginous  wall  bipolar  each  canals.  over  have s l i g h t  the  at  fertilization  the  with  the prototrichogyne  P. u m b i l i c a l i s  that  prototrichogyne  Conway and C o l e  (as  noted  monostromatic  canals,  (1959) o b s e r v e d  exists  most  spermatia  not observe  be  did  1952; Tseng and Chang, 1955;  with  whereas i n t h o s e  in  figured  association  1896  that  in  workers  flagne,  by i t ( J o f f e ,  ( B e r t h o l d , 1882 and  should  figured  the s i d e o f t h e carpogonium  1927;  with  i n one o f t h e two s p e c i e s  whereas  Several  did  It  spermatia  p r c t o t r i c h o g y n e s have been on  species  connection  carpogonium  prototrichogynes.  (1977b) p o i n t e d  there  canal-like  he d i d n o t show t h e  with  prototrichogynes  species  those  (1909) , w o r k i n g on t h e same s p e c i e s , d i d  figured  Hawkes  in  as examples),  a fine  (1882)  (although  observe carpogonia  the  cited  cited).  Berthold  protuberances  (1927)  are  spermatium  D a n g e a r d , 1927  that  t h e spermatium i s e n g u l f e d  prototrichogynes  between  and  indicated  that they  of the t h a l l u s  ended  or i n the  purpurea not  see  but without blindly i n  intercellular  143  layer.  Graves  (1969)  prototrichogynes that  the  with  Drew  the  (1956)  fertilization  of  to  nucleus  two  this  nucleus  to  in  spermatium  a  with  there fusion  spermatia  the  cell,  on  (1963)  They  figure  carpogonial  each  other  was  (1952)  male  In  the  first  yezoensis,and  the  which i s  canal connecting  the  to  a the  report  been done  by  (1969b)  on  Yabu  female nucleus  carpogonium  a  as a  reported  r e c e n t work has  and  has  fertilization  nucleus.  Magne  male  in  a  by  (1927)  which  s i n g l e large nucleus  only  a  a report  interpreted  fertilization  P.  of  and  Dangeard  for  Dangeard»s  one  wall  c a r p o g o n i u m , and  Tokida  and  cells,  a  evidence  I t c o n s i s t e d of  nucleus.  Yabu and  with  never found  cytological  cuter  is  The  contact  common and  the  carpospores.  tenera.  felt  the  what  diploid  P.  K i i t z i n g emend. A g a r d h b u t  in a  prophase i n the with  with  time.  its  cell  cell  t o be  carpogonia  adjacent  i s in contact  neighbouring assumed  that  nuclei  spermatium a t t a c h e d In  not  summarized  up  illustration  was  occasional  prototrichogynes.  (1896) o f two  canal.  capensis  phenomenon  associated  Joffe  i n P.  reported  just  i n close prior  to  fusion.  3.  R e p o r t s Of  Dangeard's fertilization Dangeard  had  f u n g a l I n f e c t i o n Of  Porphyra  (19 27)  of  canals  l e d Kunieda  observed  parasitization  c f the  His only  for  observed  basis wall  report  was  Porphyra  making  penetrations  spermatia  (1939) t o  not  a  sexual  t h a l l u s by  this i n the  suggest  forming that  fusion  an o o m y c e t b u s  statement  was  that  but  what the  fungus. he  vegetative c e l l region  had which  144  he t h o u g h t were c a u s e d by Fritsch  (1945, p.  evidence  for  this  by  Dixon  overlooked that  a  434),  sexual  Kunieda  fungus  fusion  Dangeard's  reached  the  could  be  to  the  (1977)  convincing  thalli  that  of  fertilization was  Kunieda  has  did  been  believe  and  disagreed  contents  the  spermatium  canal.  Recently  that a  the  thought  of  fertilization suggested  the;  that a  fertilization  evidence  was the  narrow  fronds,  canals  that  f i g u r e d by  fungal  of  presented.  the  infection  a  carpogcniua;  Hus  (1902) added  reports of f e r t i l i z a t i o n . the  lines  which  that  he  walls of a l l regions perpendicular  said  Berthold  He  tc  r e m i n d e d him  (1882).  No  of the  of  the  evidence  given. The  only  fungi  Porphyra t h a l l u s 1947, and  i s that  bacteria infesting  the  point  convincing  spermatium  he e x a m i n e d f o r m e d  surface  A  by  the e n t i r e  scepticism surrounding  commented  p r o v i d e any  out  Kunieda  also  for  been p o i n t e d  occurred.  concept  mistaken  however no  theory.  c a r p o g o n i u m by  Krishnamurthy  As has  d i d not  (1970, 1973)  prctotrichogyne engulfed with  a fungus,  1962; Sato,  1972;  chytrid 1973; (known  1969; et  as  the the  resulting  i n the The  Nitta,  a l . 1974; 1960;  1969).  penetrate  1975).  and  Kazama and  (Arasaki,  Sparrow,  have  been  are s p e c i e s o f Pythium  Tsuruga  Sakurai  that  (Ocmycetes)  1960;  Fuller  Fuller,  1970;  Takahashi  Sasaki  et  al.  et  a l . 1960;  In t h e c a s e  of  the  host c e l l s  and  formation  and  i n Japan),  Sasaki Sakurai,  1977)  and  fligita,  Pythium  red  pigment  infection  the f u n g a l  i s v i s i b l e t o the  crystals  unaided  eye  a  1969,  hyphae  cause a r a p i d p l a s m o l y s i s of of  the  (Arasaki,  et a l . 1966;  Arasaki  'red r o t ' disease  infection  .reported to i n f e c t  them,  (Ogata, and  does  145  not the  b e a r any r e s e m b l a n c e t o t h e r e p o r t s o f case  initial  of  stage  the  chytrid,  of infection  only  could  forming  a fertilization  canal.  Arasaki  {1960) s p e c u l a t e d  that  and  female  swarming  (1927)  statement  regarding  {1919)  of  these  of  the  but t h a t  Berthold's  reports of f e r t i l i z a t i o n .  reports  a  I t i s interesting Yendo*s  spermatium t o note  report  of  workers  and  male  may be  Dangeard*s  Furthermore, Ogata  are  that  he d i d n o t make a  (1882)  of Porphyra,  In  zoospores a t t h e i r  be m i s t a k e n f o r  infection,  a d i s c u s s i o n o f the pathology the  settled  gametes from P o r p h y r a c o n c h o c e l i s  attributable to a chytrid siiiilar  fertilization.  (1975), i n  made no m e n t i o n  that  attributable t o a fungal  infection. In view spermatia  and t h e i r  infection,  i t  fertilization examination. fertilization light,  poor  documentation  fertilization  became  apparent  a s proposed by It  was  i n Porphyra  transmission  attempt t o r e s o l v e  and  canals to  me  Berthold  therefore gardneri scanning  the problem.  that  the  were a c t u a l l y that (1882)  decided  the  a  fungal  method  merited to  reported  of  closer  t r y to  find  and t o examine i t u s i n g  the  e l e c t r o n microscopes i n an  146  Results  1.  Spermatogenesis - E l e c t r o n  Some g e n e r a l were  observed  (Figure  wall  u l t r a s t r u c t u r a l features  in  transverse  20a) c o n t a i n e d  located  pyrenoid.  that  was l a i d  transition  to  a  The n u c l e u s  small to  (Figures  developed  20c and  However, a r o u n d t h e  At  as  well  higher  fibrous vesicle.  time large  of  to  the  fibrous  at t h i s  considerably  stage  These  outside  of  I t had a d e g e n e r a t e a p p e a r a n c e , t h e r e were  several plastoglobuli. nucleus  possessed  liberation  the large  (Figure 21c), mature a  highly  a  often  nuclear fibrous  and t h e n u c l e a r  spermatium condensed  membrane. vesicle  ,  were a l s o  appeared o f , the of the  being  and  a  present,  only a few  to  and t h e  extracytcplasmic  membrane began t o d i s a p p e a r .  and l a c k e d  the  P r i o r t o spermatium  became  ( F i g u r e 22) c o n t a i n e d state,  vesicles  i n s i z e and s t r u c t u r a l  distorted  Mitochondria  final  maturation  complexity.  which  reduced  The  ( F i g u r e 21b)  was  thylakoids  the  c o n t r i b u t i n g t o the formation  Also  in  and 20d).,  chloroplast  few  seen  magnification  contents  as  undergoing the  floridean starch  v e s i c l e s were a p p a r e n t a s w e l l . their  The new  t h e immature s p e r m a t i a d i d n o t  cells  well  be  a mature spermatangium  releasing  spermatium, large  still  (Figure 21a).  fibrous  be  can  cell  centrally  located.  clearly  were p r e s e n t .  appeared  with a  was l a t e r a l l y  Ultra structurally  to give  chloroplast  vegetative  spermatangiua  much f r o m v e g e t a t i v e  division  The  cell  differ  grains  spermatogenesis  down a r o u n d a v e g e t a t i v e  20b.  was  of  section.  a stellate  Figure  chloroplast  Microscopy  a n u c l e u s which  a nuclear  membrane.  The  was i n Small  147  fibrous vesicles  were a b u n d a n t and  present.  spermatia  Most  grains. the  At  the time  spermatium;  c o n t e n t s of the from  3-5  2.  in  um  i n diameter,  examining  order to  assess  'spermatia' carpogonia similarity  - Light  floridean  by  The  1977b),  to B e r t h o l d ' s  diameter  of  the  to  this,  •spermatia* and  23c).  there with  were the  These  t h a t they  tc  this  prove  microscope.  inside  genus  examples  varied  Porphyrella of  They had  'spermatia*  suspected  of  were o n l y the In  penetration  using  found 3-5  the  um  addition  (Figures  however, i t was  without  strong  connecting  of the carpogonium  were s p e r m a t i a ;  the  fertilization  released spermatia. canals  of  a very  t h e y were i n  a t t a c h m e n t and  conclusively  the  formation  prototrichogynes  fertilization  Their specific  suggested  the  ( F i g u r e 23a).  s i z e range of the  around  um.  (1882) i l l u s t r a t i o n s  leucosticta.  wall  released spermatia  a t t a c h e d t o t h e p r o t o t r i c h o g y n e s , and in  cell  starch  a plasmamembrane and  numerous  over  no  s t a g e s of carpogonium  validity  were o b s e r v e d  Porphyra  possess  was  Microscopy  the e a r l y  attaching  not  most b e i n g 4  the  1977a,  chloroplast  surrounded  large vesicles.  While  reduced  r e l e a s e , t h e r e was  i t was  Fertilization  (Hawkes,  ih  of  did  a  the 23b  strongly impossible electron  148  3.  Fertilization  Two °f  scanning  a r e shown  spermatia  prototrichogynes. appearance,  present  on  A  the  low  the  these  the  of  demonstrates Mitochondria  m a t e r i a l can The  two  transverse  Figures  section  however, t h e  does  the  the  to  bound and  released are  also  and  is  their  presence of a  mature and  a thin  in  t o show  25b-25d.  In  locking spermatium  not  fungi.  material layer  has of  a  are  of  wall  anything  carpogonium.  been the  canals.  fertilization F i g u r e 27  are d i s t i n c t . the  wall  The the  canal  was  another  which The  fertilization  deposit of  26  to  shows in  is  Figure  transferred  carpogonium  through slight  whether  fertilization  c e n t r a l pyrenoid go  i s shewn  spermatium.  s e c t i o n s examined. cf  section  degenerate  nuclear  25c  clarify  diameter  not  the  magnification  spermatia  25b  material  and  over  already  carpogonium  i n the  spermatium t o the  section  The  of a transverse  found  non-membrane  The  prototrichogynes of  are  nuclear  24b.  Cyanophyta  reduced,  they  the  surface  shrivelled  a r e shown i n F i g u r e s  that  point  i n the  and  a  empty s p e r m a t i a  carpogenium. um  cf  seen around  from  spermatial  0.6-0.9  be  and  and  of having  micrograph  of  In  next  transferred shews  and  as w e l l .  sunken  carpogonium.  the spermatia  that  24a  visible  examples a t a h i g h e r  typical  thallus  surface.  presence  chloroplast  a  result  spermatia  Other  contents  present  a  the  thallus  of  25a.  these,  as  Figures  clearly  have  magnification  t h r o u g h one Figure  Seme  into  in  are  probably  t h e i r contents  Microscopy  e l e c t r o n m i c r o g r a p h s of t h e  Porphyra g a r d n e r i  suspected  - Electron  the plane canal;  material  in  149  the  prototrichogyne  fertilization Some  below  canal  is  spermatia  the  spermatium  that  have  transferred  r e t a i n what a p p e a r s t o be  (Figures  28a  thylakoids the  nuclear  plugged This of  28b).  and  by  the  28c  apex o f  and  a  apex o f  the  Following  fertilization deposited  the  go  chloroplast  a  nuclear  chloroplast  few  the  distorted  transfer  canal  around  Carpogonia  29),  details  is  of  rapidly  i t (Figure  28b).  i n the  with  which  but  of  the  apex  and  of  with The  first  c y t o k i n e s i s to 30b.  The  the  plane  canal.  In  the  mature s p e r m a t i u m .  Another  28d.  I t would from  the  nuclear  t o be  unable  division  of  to  material. highly  observed  l a c k of a  carpogonial  that  spermatium  were  apparent  give  appear  appeared  n u c l e u s I was the  empty  typical  material the  an  reduced c h l o r o p l a s t  what  nuclear  fusion  and  30a  fertilization  spermatial  contained  spermatial  fusion  nucleus,  i n the  the  because  level.  visible.  the  through  Onfortunatly  have been t r a n s f e r r e d  ultrastructural  Figures  through  spermatial  membrane a r o u n d  section  carpogonium.  found  carpogonium along  (Figure  the  i s shown i n F i g u r e  these chloroplasts  condensed  transverse  carpogonium i s a s m a l l  example o f t h i s  in  the  material  is  o f s e c t i o n d o e s not  the  a  carpogonium.  spermatium  the  remains of  T h e s e c h l o r o p l a s t s have a  material, wall  their  a p p e a r s t o o r i g i n a t e from dictyosome a c t i v i t y  Figure  of  the  some p l a s t o g l o b u l i .  new  that  present.  material  and  indicates  nuclear  to  follow  nucleus at the  an  presumed  2 c a r p o s p o r e s , i s shown  fertilization  canal  is  still  150  4.  Chromosome  Evidence the  Counts  that a fusion of spermatial nuclear  carpogonial nucleus  chromosome c o u n t s . cells  of  the  chromosomes  diploid and  Hawkes  foliose  (n=4)  carpospores  does  and  occur  been  (1977b) r e p o r t e d thallus  and  ( F i g u r e s 31a vegetative  chromosome number  has  obtained  with from  that the vegetative  the  spermatia  and 31b) i  cells  material  In  have  both  of the c o n c h o c e l i s  2n=8 has been o b t a i n e d  4  the  phase a  ( F i g u r e s 31c,d  31e,f).  5.  Feulgen  In the  Staining  an a t t e m p t t o c l a r i f y  spermatial nuclear  the d e t a i l s of  of  using  Initially  Feulgen  transfer  m a t e r i a l t o the carpogonium,  sections of carpogonial regions the  the  technique-.  stained  t h e s t a i n was u s e d on  undergone  maximum number o f d i v i s i o n s , b u t t h e n u c l e u s i n  condensing released which  of  agree with  can  fertilization  canal  nucleus likewise  i s  seen  observations  present.,  containing  attached  in  a  Figures  to  3 3c  state. 32b.  nuclear These  The The  material  observations  condensed  nuclear  the prototrichogyne. the  s t a t e and a and 33d  has  of spermatogenesis.  I n F i g u r e 33b  diffuse  which  i n Figure  condensed  (Figure 32c).  c a n a l i s present. is  interphase  complete  have  33a a s p e r m a t i u m be  spermatangium  diffuse  a dense p u r p l e  material is  in a  the u l t r a s t r u c t u r a l  Figure  material  nuclear  the  spermatia  stains  In  is still  a  were  Figure  most s p e r m a t i a  shows  thallus  transverse  the spermatia. the  32a  the  of  show  No  spermatial  fertilization condensed  151  spermatial  nuclear  material  present.  The c o r r e c t t i m e  known.  It  may  be  and  fertilization  sequence f o r  that  the  Figures  spermatial  canals  33b-d  are  i s not  nuclear  material  deccndenses i n order  t o c o d e f o r an enzyme r e q u i r e d  fertilization  and t h e n r e - c o n d e n s e s f o r t h e t r a n s f e r  the  canal,  carpogonium.  fertilization in  canal  the released  material being  Another  explanation  enzymes a r e p r e s e n t  s p e r m a t i u m , and  remains  followed  condensed by  that  until  would  be  (or at l e a s t the  that  and  to the  coded f o r )  spermatial  the canal  de-condensation  t c make t h e  nuclear  i s complete,  transfer  to  this the  carpogonium. Once t r a n s f e r has o c c u r r e d shews mass of  up i n t h e c a r p o g o n i u m a s a s m a l l (Figure  33e).  33g).  (Figures On fact  not with  material  densely s t a i n i n g  purple  carpogonial  up  to  4  nucleus  or  5  (Figures 33f  have  been  observed  33h and 3 3 i ) . the b a s i s  that  of the d i p l o i d  dense  carpogonial  spermatial  large  nucleus,  suspected  33j).  first  nuclear  and t h a t  division  33k).  followed  by  number,  this fusion;  fusion  The  cf  divisons  to  the  fusion and  (Figure give  them  fuses  degenerate. however,  4  331) and  the  pyrenoid to give 8  is  I t has  been  ultrastructural nucleus  with  carpogonia  n u c l e i have f r e q u e n t l y  chloroplast  cytokinesis  and  masses a r e n o t f o u n d i n  the rest  As h a s been shown a t an  (Figure  Subseguent  chromosome  i t i s assumed t h a t one o f  been p o s s i b l e t c o b s e r v e  (Figure the  t o the  Occasionally  subseguent stages, the  nuclear  I t i s n o t uncommon t o s e e from one t o t h r e e  these i n a d d i t i o n  and  the spermatial  seen  level,  periclinal  then  divide  2 carpospores.  carpospores  per  152  carposporangium division  are  sequence  involved  Porghyra g a r d n e r i (see  shown i n F i g u r e s  has  in  33m-o.  A summary o f  the  formation  by  carpospore  been r e p o r t e d  previously  (Hawkes, 1977b)  Part I I ) .  Discussion  1.  Spermatogenesis  The are for  main f e a t u r e s  essentially  the  of spermatogenesis i n same as t h o s e r e p o r t e d  Bangia fuscopurpurea  respects  t o the  Smithgra  naiadum.  indicated  i f the  Porphyra  report  (Dillwyn)  of McBride  L y n g b y e , and (1972)  Unfortunatly nuclear  by  McDonald  membrane r e m a i n e d  (1972)  s i m i l a r i n some  for the  neither  gardneri  of  spermatia these  intact  of  workers  i n the  mature  spermatium. There  are  some  spermatogenesis  in  R2I£hlL\sk g a r d n e r i .  The  have  in  vesicles, spermatium complexity  striking the  most c h a r a c t e r i s t i c  the  is  and  r e l e a s e of the  liberation. in  the  production  The  have i n common., K u g r e n s  of  possible explanation  (1974)  f o r the  fibrous to  they  a  general  i n s e v e r a l qroups.  Another  absence  Kuqrens  large  in chloroplast  (1974) n o t e s t h a t t h i s  qeneral  they  spermatium i s a n o t h e r f e a t u r e  sinilarity  mature s p e r m a t i a .  and  that  and  algae  the  small  in  size  phenomenon i n male gametes o f is  feature  and  large fibrous v e s i c l e prior  reduction  mature  between  Florideophycidae  common the  similarities  has  reduction  of  starch  pointed  is  qrains i n out  that  the one  i n chloroplast structure  153  and  storage  developing  product  i s  to  prevent  on i t s own, t h e r e b y  Most  workers  report  bypassing  that  the  spermatium i n t h e F l o r i d e o p h y c i d a e (Grubb,  1925;  Peyriere, Scctt in  Fritsch,  1974; Young,  and D i x o n  Ptilota  the  male  sexual  nucleus  gamete  reproduction. of  the  c o n t a i n s condensed  1977), a s i s t h e  case  in  spermatia,  but  judging  from  1974) does n o t c o n t a i n c o n d e n s e d of  of Porphyra  previously  been r e p o r t e d  several and  explain  at  Florideophycidae  spermatium  the  this  is  little  nuclear  membrane i s p a r t i c u l a r l y  fixation an  procedure  intact  released  nuclear  spermatia  the  that  For  the  level  Dixon  at  (1973b).  i n the breakdown  may be t h a t t h e this  stage  was n o t p r e s e r v e d i n those  examined  in  i td i f f i c u l t to  has been r e p o r t e d , to  cases  of  by t h e where  perhaps c i d e r see  i f the  disappear.  Florideophycidae  (Peyriere,  T h i s has  1945, p . 596)  membrane  sensitive  have been  only dictyosome involvement vesicles  chromatin.  disappear  The c a s e  Alternatively,  membrane  membrane does e v e n t u a l l y  nuclear  Rhodophyta.  used.  and  would  i t therefore  should  microscope  1925; F r i t s c h ,  membrane  mitosis  and  light  b e c a u s e he f o u n d  during  development  in  Janczewskia  membrane i n t h e mature  l e v e l by S c o t t  nuclear  when t h e r e  the  (Grubb,  (1974) g u e s t i o n e d why  the nuclear  in  g a r d n e r i i s r a t h e r unexpected.  a t the u l t r a s t r u c t u r a l  Kugrens  nucleus  condition  micrographs,  spermatia  disappearance  The  their  i s present.  spermatium  chromatin  P. g a r d n e r i .  condensed chromatin  The  mature  1945, p. 596; K u g r e n s and l e s t , 1972;  (1973b) were u n c e r t a i n o f t h e n u c l e a r  (Kugrens,  from  spermatia,  i n the 1971;  many w o r k e r s r e p o r t  formation  Scott  and  of  the  Dixon,  fibrous 1973b;  154  Simon-Bichard-Breaud, Kugrens  (1974) and  involved fibrous  1972a);  Young  i n the early  however, K u g r e n s and  (1977) have i n d i c a t e d  West  that the  from  EB  EB  stages i n the s p e c i e s they s t u d i e d .  v e s i c l e s i n t h e s p e r m a t i a o f P. g a r d n e r i may  derived  (1972),  as  few  dictyosomes  were  is The  be  mostly  observed  during  spermatogenesis. R e p o r t s on t h e c h e m i c a l n a t u r e and v e s i c l e s have been summarized appear  to c o n t a i n a  analysis of  is  the  pressure  spermatangial wall Likewise,  the  attachment been in  other  involved  snyderae  widespread red  algal  IV),  unpublished (Kugrens Taylor Ptilota  and  in  i n spermatium  spores  such  of  observation), 1973,  (Wetherbee hypnoides  ( S c o t t and  must have an i m p o r t a n t  in  (Fetter,  1972) Porphyra  Levringiella  1974)  and  increase  and  Wynne, Dixon,  function.  the  release. spermatium  and  fibrous the  have  dispersal  vesicles  in  monospores  of  Cole, and  in  1977).  1971),  Porphyra g a r d n e r i  nereocystis and  and  (Hawkes,  Ervthrocystis  Polysiphonia 1973)  of  strands  release  as  function  spermatium aid  They  critical  rupturing  and  (McDonald,  carpospores  an  R e c e n t l y , mucilage  (McBride  West,  may  (Farlow) Abbott  Porphyropsis coccinea Part  the  fibrous  (1973b).  suggested  cause  occurrence of s i m i l a r  S m i t h o r a naiadum  see  in  mucopolysaccharide  Dixon  One  may  and t h e r e b y a i d i n g  t o be  Tiffaniella  they  resulting  to the t r i c h o g y n e .  shown  The  i s that  of the  a l t h o u g h more  to confirm t h i s .  vesicles  osmotic  and  mucopolysaccharide,  reguired  the f i b r o u s  by S c o t t  function  novae-anqliae  tetraspores  1973a) i n d i c a t e s  that  of they  155  2.  Fertilization  This  is  the  first  conclusive  occurrence of a sexual fusion except in  f o r Hagne's  in a  documentation  species  (1960) e x c e l l e n t  of  of  the  Porphyra.  and  work on s e x u a l r e p r o d u c t i o n  Bhodochaete p a r v u l a T h u r e t i t i s the o n l y u n e g u i v o c a l r e p o r t  of  fertilization  classical  interpretation  reproduction the  i n the Bangiophycidae.  in  Porphyra  observations  Dangeard  by  (1927),  appear  leucosticta  Magne  (1963),  be  basically  ( 1 9 2 1 ) , Dangeard divison  of  the  (1909),  of  Chang  Similarly, (1921),  (1955),  (1967b)  the  sexual  Ishikawa  and  Migita  ,  Kurogi  and  Yabu  process i n other Porphyra s p e c i e s  correct.  (1927) and  (1882) i s correct.  ( 1 9 5 2 ) , Tseng  o f the f e r t i l i z a t i o n to  Berthold  Bosenvinge  ( 1 9 6 1 ) , Yabu and T o k i d a (1969b)  by  i t confirms that  Tseng  carpogonium  The and  reports  Chang  being  of  Ishikawa  (1955) o f t h e  meiotic  are  first  probably  incorrect. The Derbes Suto  mechanisms o f and  Solier  (1963) and  Kunieda  to  s e v e r a l o r even the  (cited  for  Porphyra  from  Drew,  (1939) a r e p r o b a b l y i n c o r r e c t . fertilization  produce  carpospores  many v e g e t a t i v e c e l l  and  the  division  by  1956), Dixon*s of  the  are s e p a r a t e d i n time divisions  i s likewise  by not  case. The  (1973,  and  chromosome  reports  by  Krishnamurthy  i n p r e s s ) and number  phase of Porphyra the  proposed  (1856), Koschtsug  (1973) s u g g e s t i o n t h a t carpogonium  fertilization  tremendous  in  Mumford and the  (1959), Cole  (1977) o f  vegetative c e l l s  are not n e c e s s a r i l y adaptability  Conway  by  a  Cole  haploid  o f the conchocelis  incorrect.  exhibited  and  In  view  the Porphyra  of life  156  history  i t i s possible that carpospore  parthenogenetically. Nichols  (1970)  conchocelis  of  of  development o f formation not  Similarly, a  Bangia  fertilization  canals.  apcgamously.  Kurogi  f o r which  Drew  and  (1972)  that  the  others of  seme c a r p o s p o r e s  are the r e s u l t  this  fusions  point are  fertilization  canals  have a l s o been  haplcid  other  conchocelis division  phase  takes  he  could  of sexual  not  out there  a r e some canals.  (1973)  carpospores fusions.  have  which a r e  The  case  parthenogenetically Further  find  developed  fertilization  of  the  may  whereas  investigation  C e r t a i n l y i n Porphyra g a r d n e r i the  by  there  often  Rosenvinge  beinq  several  Multi-penetrations  (1909)  and  Danqeard  umbilicalis. explanation  number is  in  that  of the f e r t i l i z e d  T s e n g and Chang  observed  o f p l a n t s which d i d  i n t o one c a r p o q o n i u m .  possible  chromosome  number  the  parthenogenetic  (1959) and D i x o n  abundant,  reported  (1927) f o r P o r p h y r a The  observed  occur  in  carpospores  of f e r t i l i z a t i o n .  extremely  a  pointed  c a n form  i s required.  number  (1909)  which  has  could  by Sommerfeld and  to  He s u g g e s t e d t h e  produced a r e a l l t h e product that  in  he has n e v e r  doubt  due  i n a population  (195tta), K r i s h n a m u r t h y  expressed  be  be  Rosenvinge  of carposporangia spermatia  the report chromosome  may  carpogonia;  produce  species  haploid  formation  f o r the occurrence  the  vegetative  meiosis  occurs  cells during  of  i s no e v i d e n c e  the  the f i r s t  c a r p o g o n i u m as has been r e p o r t e d  ( 1 9 5 5 ) ; however, t h e r e  of a  that  by this  place. Although  the s i t e  of meiosis  been i n v e s t i g a t e d y e t i t p r o b a b l y  i n Porphyra  g a r d n e r i has n o t  occurs  the  in  conchospore  157  branch  as  Migita  has  view o f t h e  fertilization  has  and  cells  genus  is  technigues Porphyra  future the  The  as  in  may  have  other  by  occuring  According  be  Porphyra  of  not to  in  contact.  developed  which  microscope  level,  carpospores  sexual The  reproduction application  in  this  species.  I t i s expected  in of  study  in  of  demonstrating  h i s t o r y found i n the (1967b)  or  reported,  helpful  Janczewski  i n the  in his  show  that that  genus i s t h e  excellent  work  Fritsch  spermatia  and  as  i n Porphyra  a result  of  spermatium r e a c h  (except  the  certain  Hemaliales) the  transfers i t s contents  from  which  prototrichogyne  and  a l s o notes that  the  carpogonium in  Such a  carpogcnius  wall d i s s o l u t i o n i n  differs  and  the  (1945, p. 597)  g a r d n e r i i n which the chloroplast  enigma.  gardneri.  (1945, p. 592)  Florideophycidae  This  of  same s p o r a n g i u m r e m a i n s an  Flcrideophycidae  the  (1873)  m i s i n t e r p r e t a t i o n by J a n c z e w s k i .  observed  i n the  trichogynes  Porphyra  for  phase h a v e been  used  by  (1974).  Porphyra s p e c i e s w i l l  Migita  lack chloroplasts., Fritsch spermatium  species  jezcejisis.  phenomenon was  the  been  cytological l i f e  have b e e n a  trichogyne  Kito  light  occurrence.  should  report  carpospores  at the  to suggest that  common  t h a t o u t l i n e d by  on P o r p h y r a  (1968) and  Porphyra  chromosome numbers f o r t h e  i n v e s t i g a t i o n s of other  same a s  This  a  gardneri  typical  other  of t h e c o n c h o c e l i s  such  fertilization  Magne  been r e p o r t e d  would seem r e a s o n a b l e  the  for  l a r g e number c f P o r p h y r a s p e c i e s  i n which d i p l o i d  vegetative it  reported  (1967b) G i r a u d  In  and  been  the  the  region  situation  possesses  a  contents  of  via a f e r t i l i z a t i o n  to  in well the  canal.  158  A d e t a i l e d e x a m i n a t i o n and the  ultrastructural  level  Flcrideophycidae Nemaliales)  and  i n suspected  (for other  example;  members o f  interesting  insight  reproductive  system i n these  Although of the  history  of  need o f  further  undergo  from  the  into  this  question  wall  is  w h i c h was  (1945, p. 597) spermatial Such a  once  not  observed  that  The  i t  has  answered  reproduction  does  attachment  being  most w o r k e r s Magne,  1952;  to the  in  (Grubb,  a  the  in  major  the  life  the  in  this  study  possesses a  liberation. and  P.  the  thin  Fritsch  Nemalion  the  the  trichcgyne.  gardneri  but  spermatial  the the  nuclear  prototrichogyne  needs t o  argue  the  against  1925;  Krishnamurthy,  state  Fritsch, 1959;  to of  once i t e n t e r s  prophase  in  spermatium  time c f attachment  divide  of  b e c a u s e i t would seem t o nucleus  in  female  s e v e r a l areas  i t contacts  does  decondensing  the  Bhodophyta.  spermatium  divides  explained  Drew, 1951;  cf  r e p o r t s t h a t i n Batrachospermum  after  by  provide  at t h e t i m e o f  material  assumed  B a n g i o p h y c i d a e may  observed  attached  present  exists  spermatial  other  changes  change  the  phenomenon was  carpogonium.  and  i t p o i n t s out  What  One  nucleus  possibility  the  groups of  gardneri,  that not  p r i m i t i v e members c f  time o f r e l e a s e to the  prototrichogyne? gardneri  two  of sexual  study.  at  evolution  investigation  Porphyra  of f e r t i l i z a t i o n  Acrochaetium  the  the  occurrence  the  comparison  nature  as has  1945,  be  been  p.  Mumford and  596; Cole,  1977). Histochemistry vegetative carpogonium.  cells  may  help  undergo The  to  clarify  in  the  cuticle  of  changes  that  transformation vegetative  to  cells  the a of  159  Porphyra  u m b i l i c a l i s has  Craigie, contain  1969;  region cf  percentage  investigation may  recognition With  1963,  i n Fucoids modified  microscopy  1964)  i t  material  in  level.  With such  a  The Porphyra  may  An  and  be  be  and  and to  glycoprotein.  A  prototrichogyne  mechanisms o f  the  approach  the l i n e s  along  here.  procedures  t h a t can  be  cf  the  egg-sperm  used  with  and  Brinkley,  p o s s i b l e t o f o l l o w the  speraatial  i t may  be  Bryan  specificity  on  appropriate  carpogonium  technique  (Hanic found  i n the  ( J u r a n d e t a l . 1958;  the  1974)  Evans e t a l . (1977)  Feulgen  of the fusion known  protein  into  by  should  nuclear  formation  out  Williamson,  wall chemistry  attachment.  work c a r r i e d  electron  of  of  provide insight  spermatium  recent  Gunawardena and  a high  similar  been s t u d i e d b i o c h e m i c a l l y  at  an  ultrastructural  p o s s i b l e to observe  the  nucleus.  morphological  g a r d n e r i i s summarized  and  cytological l i f e  in Figure  34.  history  of  160  D.  E v i d e n c e Of  Sexual  Porphyra  Reproduction In Porphyra  nereocystis  and  thuretii  Introduction Following reproduction of  P.  the in  Porphyra  nereocystis  reproduction  detailed  and  in their  investigation  gardneri  sexual  I made a p r e l i m i n a r y  P. t h u r e t i i  life  of  for  evidence  of  survey sexual  histories.  Results  1«  Porphyra  The  nereocystis  spermatium  ultrastructurally the  chloroplast small  and  attached canals  the  near with  the  Porphyra nereocvstjs  the  Figure  s p e r m a t i a of 35  thallus  shows  a  gardneri  spermatium  margin.,  plastcglobuli,  P.  is  Note  c e n t r a l nucleus,  and in  a  the  reduced  and  abundant  large fibrous vesicles.  At t h e  £•  s i m i l a r to  Flcrideophycidae.  spermatangium  of  light  to  the  were p r e s e n t  nereocystis  is  fertilization  attachment  of  p e r h a p s due  to the  microscope  level  wall  the  over  (Figures  36a  spermatia  carpogonia  and  36b).  t h i c k e r t h a n i n P. canals  are  longer.  spermatia i s not absence o f  and  The  gardneri Note  as s p e c i f i c  were  as  prototrichogynes.  observed  fertilization cell  and  as  wall a  of  result  also  that  the  i n P.  gardneri.  An  electron  161  micrograph  of  associated  a  fertilization  Evidence chromosome  that  thallus  whereas  the  chromosomes,  c a n a l i s shown i n F i g u r e  The s p e r m a t i a  carposporangia  attached  at  microscope l e v e l  obtain  n=3  ( F i g u r e s 37a  conchocelis  to the s l i g h t  and  phase  from of t h e  and 3 7 b ) , have  6  37f,g).  prototrichogyne  associated f e r t i l i z a t i o n  spermatia I  h a s been o b t a i n e d  thuretii  carpogonium  Unfortunatly  and  36c.  and v e g e t a t i v e c e l l s  2n=6 ( F i g u r e s 37d,e and  Spermatia  The  occurs  have 3 chromosomes,  Porphyra  the l i g h t  s e c t i o n t h r o u g h a s p e r m a t i u m and  fertilization  counts.  foliose  2*  transverse  have  have not  2  canals  were  of  the  observed  ( F i g u r e s 36d and 3 6 e ) . chromosomes,  had  counts f o r the carpospores  sufficient  n=2  (Figure 37c).  m a t e r i a l or time to  o r t h e c o n c h o c e l i s phase.  162  Discussion T h i s p r e l i m i n a r y i n v e s t i g a t i o n o f Porphyra n e r e o c y s t i s and P. t h u r e t i i suggests occurrence that  of  that  i n their l i f e other  reproduction  workers  sexual histories. strongly  reproduction  is  a  common  By evidence combined suggests  that  with  sexual  i n the genus Porphyra i s a widespread phenomenon.  163  T a b l e V I I I . The s p e c i e s of P o r p h y r a f o r which d i p l o i d chromosome numbers have been r e p o r t e d f o r t h e c a r p o s p o r e s o r v e g e t a t i v e c e l l s of the c o n c h o c e l i s phase.  Taxcn -1. D i p l c i d  Reference  Carpospores  P°££il£a a b b o t t a e K r i s h n a m u r t h y £• d e n t a t a K j e l l m a n P. g a r d n e r i (Smith e t H o l l e n t e r g ) Hawkes £• k a n a k a e n s i s Mumford P. k a t a d a i M i u r a P. k i n o s i t a i (Yamada e t Tanaka) Fukuhara P. k u n i e d a i K u r o g i P. l i n e a r i s G r e v i l l e  P. m i n i a t a  £• P. P. P. P. £• £•  (C. Agardh) C.  Agardh  roo££ejttsis Oh mi n e r e o c y s t i s Anderson o k a m u r a i Oeda o n o i Oeda £ § £ f g r a t a J . Agardh fisejii^linearis Ueda £U££JJ£e§. (Both) C. A g a r d h  P. s c h i z p p h y l l a H o l l e n b e r g i n S m i t h and H o l l e n b e r g P. s e r j a t a K j e l l man £• t e n e r a K j e l l m a n  £• JE»  u m b i l i c a l i s (Linnaeus) J . Agardh y e z o e n s i s Oeda  Mumford and C o l e (1977) Yabu (1971) Hawkes (1977a, 1977b, and t h i s t h e s i s ) Mumford and C o l e (1977) K i t o (1966) - a s P. s p . Yabu (1972) Yabu (1972) Yabu (1971) Magne (1952) K i t o e t a l . (1967) - as £• u m b i l i c a i i s f. l i n e a r i s ( G r e v i l l e ) Harvey K i t o e t a l . (1967) - a s P. a m p l i s s i m a ( K j e l l m a n ) S e t c h e l l e t Hus Yabu (1970) - a s P. a m p l i s s i m a K i t o e t a l . (1971) K i t o e t a l . (1967) Hawkes ( t h i s t h e s i s ) Yabu (1969a) Yabu (1972) Mumford and C o l e (1977) Yabu (1969a) G i r a u d and Magne (1968) as P. u m b i l i c a l i s var. l a c i n i a t a (Lightfoot) Thuret K i t o e t a l . (1971) Mumford (1975) Mumford and C o l e (1977) Yabu (1969a) Fujiyama et a l . (cited from Yabu, 1975) K i t o (1968) Yabu (1969b) K i t o e t al.„ (1971) Yabu and T o k i d a M i g i t a (1967b) Yabu (1969a)  (1963)  16  P. s p p . # 1 and 2 Diploid Porphyra  Yabu  (1971)  Conchocelis gardneri  E* J s a n a k a e n s i s P. m i n i a t a P. p e r f o r a t a P. p s e u d o l i n e a r i s !• p u r p u r e a P. s c h i z o p h y l l a P. t e n e r a P. .. y e z o e n s i s  Hasrkes (1 977a, and t h i s thesis) Mumford a nd C o l e (1977) K i t o e t a.1. (1971) Mumford a nd C o l e (1977) K i t o (197 <i) G i r a u d an d Magne (1968) a s P. um fcilicalis var. l a ci n i a t a Mumford a nd C o l e (1977) K i t o (197 il) K i t o (196 7) M i g i t a (1 967b) K i t o (197 <*)  165  Fiqure  16a.  T r a n s v e r s e s e c t i o n o f the b l a d e m a r g i n o f Lamj.naria s e t c h e l l i i and t h e b a s e c f £2I£hyi.a g a r d n e r i . The r h i z o i d a l f i l a m e n t s be s e e n p e n e t r a t i n q i n t o t h e m e d u l l a o f t h e Laminaria blade.  Fiqure  16b.  Closeup o f medulla.  Figure  16c.  Transverse s e c t i o n thrcuqh a N e r e o c y s t i s s t i p e and t h e b a s e o f a P o r p h y r a n e r e o c y s t i s a t t a c h e d to i t .  Fiqure  16d.  Closeup of disc.  Scale  bar  List  =  Of  100  um  on  the  the  rhizoidal  penetration  c e n t r a l reqion  of  the  into  can  the  attachment  a l l Figures.  Abbreviations  Dsed On  Figures  For  Fart  IV.  C a r p o g o n i a l n u c l e u s (cn) ; c h l o r o p l a s t ( C , c ) j c e l l w a l l (CW, cw); d i c t y o s c a e (D, d ) ; f l o r i d e a n s t a r c h (FS, f s ) ; l a r g e f i b r o u s v e s i c l e ( L F V I f v ) ; n e d u l l a (ME); m i t o c h o n d r i o n (MI, J e f e o c y s J i s (NF) ; n u c l e u s (N, n) ; P o r p h y r a (PO) ; p y r e n o i d (P, p) ; s m a l l f i b r o u s v e s i c l e (SFV, s f v ) ; s p e r a a t i a l n u c l e a r m a t e r i a l ( a ) ; spermatium (S). f  166  167  Figure  17.  Tranverse s e c t i o n of Pprchyra g a r d n e r i t h a l l u s , t h r o u g h a young monosporangium. Note d i c t y o s o m e s , s m a l l f i b r o u s v e s i c l e s and l a y e r o f f i b r o u s m a t e r i a l b e i n g formed on t h e o u t s i d e o f t h e plasma membrane.  168  169  Figure  18.  Transverse s e c t i o n of Eorphyra g a r d n e r i t h a l l u s t h r o u g h a monosporangium w h i c h h a s d e v e l o p e d beyond t h e s t a g e shown i n F i g u r e 17., S m a l l f i b r o u s v e s i c l e s a r e abundant and l a r g e fibrous v e s i c l e s a r e b e g i n n i n g t o form.  170  171  Figure  19a.  Dictyosome a c t i v i t y vesicles.  producing  small f i b r o u s  Figure  19b.  C l o s e u p of a monosporangial w a l l showing t h e depositon of f i b r o u s m a t e r i a l along i t s inner s u r f a c e (arrow) .  Figure  19c.  Honosporangium i n t h e f i n a l s t a g e s o f d i f f e r e n t i a t i o n p r i o r t o monospore l i b e r a t i o n , B o t h s m a l l and l a r g e f i b r o u s v e s i c l e s a r e abundant.  172  173  Z21£lJHS  gardneri - spermatogenesis, e l e c t r o n (transverse sections)  Figure  20a.  Vegetative c e l l  Figure  20b.  The f i r s t d i v i s i o n o f t h e s p e r m a t a n g i u m i s n e a r l y complete. Mote t h e new w a l l l a y e r (arrow) t h a t has been l a i d down i n t h e t r a n s i t i o n from a vegetative c e l l .  Figures  Scale  20c-d.  t a r = 5 um  later  of the f c l i o s e  microscopy  division  s t a g e s o f the  on a l l F i g u r e s .  thallus.  spermatangium.  174  175  Porphyra  gardneri - spermatogenesis, e l e c t r o n (transverse sections)  microscopy  Figure  21a.  F i n a l d i v i s i o n s i n the formation of a nature spermatangium. Note t h e p r o d u c t i o n o f l a r g e fibrous vesicles.  Figure  21b.  Spermatium from a mature s p e r m a t a n g i u m . It c o n t a i n s a nucleus, mitochondria, reduced c h l o r o p l a s t , s e v e r a l s m a l l and one l a r g e f i b r o u s vesicle. Note s m a l l f i b r o u s v e s i c l e e m p t y i n g i t s c o n t e n t s i n t o the l a r g e f i b r o u s v e s i c l e (arrow).  Figure  21c.  Two s p e r m a t i a i n a s p e r m a t a n g i u m n e a r t h e r e l e a s i n g margin. The l a r g e f i b r o u s v e s i c l e j u s t become e x t r a c y t o p l a s m i c .  Scale  bar  =  1 um  i n Figures  21b-c  has  176  177  Figure  22.  Mature spermatium j u s t p r i o r t o r e l e a s e . Note t h e n u c l e u s w i t h h i g h l y c o n d e n s e d c h r o m a t i n and no n u c l e a r membrane, r e d u c e d c h l o r o p l a s t w i t h numerous p l a s t o g l o b u l i , m i t o c h o n d r i o n and abundant s m a l l f i b r o u s v e s i c l e s .  178  179  Porphyra g a r d n e r i - f e r t i l i z a t i o n , (transverse sections)  microscopy  Figure  23a.  Spermatium a t t a c h e d carpogonium.  Figure  23b.  Two s p e r m a t i a w i t h f e r t i l i z a t i o n i n t c the carpogonium.  Figure  23c.  Spermatia a t t a c h e d t o both p r o t o t r i c h o g y n e s f e r t i l i z a t i o n canals).,  Scale  bar =  10  um  on  a l l Figures.  to the  light  prototrichogyne canals  of  the  going (note  180  181  Figures  24a-b. Two s c a n n i n g e l e c t r o n m i c r o g r a p h s o f t h e t h a l l u s s u r f a c e o f P o r p h y r a g a r d n e r i showing s e v e r a l spermatia attached t o the prototrichogynes.  182  183  Porphv.ra g a r d n e r i - f e r t i l i z a t i o n , (transverse sections) Figure  25a.  Figures  Scale  electron  microscopy  Low m a g n i f i c a t i o n view o f a s p e r m a t i u m to t h e p r o t o t r i c h o g y n e .  attached  25b-d. Three examples c f spermatia a t t a c h e d t o t h e prototrichogyne p r i o r to the formation of the f e r t i l i z a t i o n canal. They c o n t a i n a c h a r a c t e r i s t i c reduced c h l o r o p l a s t , mitochondria and non-membrane bound n u c l e a r m a t e r i a l .  b a r = 2 um on F i g u r e s  25b-d  184  185  Porphyra Figure  26.  gardneri  -  fertilization  T r a n s v e r s e s e c t i o n t h r c u g h a c a r p o g o n i u m and two s p e r m a t i a w h i c h have t r a n s f e r r e d t h e i r n u c l e a r m a t e r i a l dcwn the f e r t i l i z a t i o n c a n a l i n t o t h e carpogonium.  186  187  Porphyra Figure  27.  gardneri -  fertilization  T r a n s v e r s e s e c t i o n through a carpogonium showing b o t h p r o t o t r i c h o g y n e s , c e n t r a l p y r e n o i d and l a t e r a l nucleus. The p r e s e n c e o f a s m a l l d e p o s i t of w a l l m a t e r i a l (arrow) i n the prototrichogyne below t h e s p e r m a t i u m i n d i c a t e s t h a t a f e r t i l i z a t i o n c a n a l has been f o r m e d , a l t h o u g h i t i s not v i s i b l e i n t h e plane of s e c t i o n . S u s p e c t e d C y a n c p h y t a a r e a l s o v i s i b l e on t h e t h a l l u s s u r f a c e (double arrow).  188  189  Porphyra g a r d n e r i - f e r t i l i z a t i o n (transverse sections)  Figures  28a-fa. Two s p e r m a t i a which have t r a n s f e r r e d t h e i r n u c l e a r m a t e r i a l down t h e f e r t i l i z a t i o n c a n a l i n t o t h e c a r p o g o n i u m , but w h i c h s t i l l have t h e r e m a i n s o f a c h l o r o p l a s t l e f t i n them.  Figures  28c-d., P r o t o t r i c h o g y n e r e g i o n o f t h e c a r p o g o n i u m w i t h a r e d u c e d c h l o r o p l a s t which i s t h o u g h t t o h a v e come from t h e s p e r m a t i u m d u r i n g t h e t r a n s f e r o f nuclear material.  Scale  car =  1 um on a l l F i g u r e s  190  191  Porphyra Figure  29.  gardneri -  fertilization  T r a n s v e r s e s e c t i o n t h r o u g h a c a r p o g o n i u m which has t h e r e m a i n s o f a s p e r m a t i u m a t t a c h e d t o t h e p r o t o t r i c h o g y n e , and what may be t h e h i g h l y condensed s p e r m a t i a l n u c l e a r m a t e r i a l (arrow).  192  Por p h y r a g a r d n e r i carposporogenesis (transverse sections) Figure  30a.  F e r t i l i z e d c a r p o g o n i u m showing t h e two d a u g h t e r n u c l e i r e s u l t i n g frcm the f i r s t d i v i s i o n of the fusion nucleus. The f e r t i l i z a t i o n c a n a l i s s t i l l p r e s e n t a l t h o u g h t h e s p e r m a t i u m has disintegrated.  Figure  30b.  The f i r s t d i v i s i o n o f t h e c a r p o s p o r a n g i u m i s p e r i c l i n a l and p r o d u c e s 2 c a r p o s p o r e s (note the f e r t i l i z a t i o n c a n a l s t i l l present i n the w a l l ) .  Scale  bar  =  5 um  on  a l l Figures  194  195  P o r p h y r a g a r d n e r i - chromosome c o u n t s Figure  31a.  Spermatium,  Figure  31b.  Vegetative  n=4. cell  Figures  31c-d.  Figures  31e-f. Vegetative 2n=8.  Scale  Carpospore,  of t h e f o l i o s e  thallus,  n=4.  2n=8. cell  b a r = 5 um on a l l F i g u r e s .  of the conchocelis  phase,  96T  197  Porphyra g a r d n e r i - Feulgen s t a i n i n g {transverse sections)  Figure  32a.  M a t u r e s p e r m a t a n g i u m ; most s p e r m a t i a s t i l l have n u c l e i i n a d i f f u s e i n t e r p h a s e s t a t e (arrow).  Figure  32b.  Mature s p e r m a t a n g i u m r i g h t on t h e r e l e a s i n g margin. The n u c l e a r m a t e r i a l i s h i g h l y c o n d e n s e d (arrow).  Figure  32c.  Released material  Scale  bar =  10 um  s p e r m a t i a with (dark d o t s ) .  on a l l F i g u r e s .  h i g h l y condensed  nuclear  198  %  ****  m  199  Porphyra g a r d n e r i - Feulgen s t a i n i n g (transverse sections) Figure  33a.  C a r p o q o n i u m and a t t a c h e d s p e r m a t i u m w h i c h c o n t a i n s condensed n u c l e a r m a t e r i a l . No f e r t i l i z a t i o n canal i s present.  Figure  33i>,  C a r p o g o n i u m and c a r p o g o n i a l n u c l e u s l a t e r a l l y located. The a t t a c h e d s p e r m a t i u m c o n t a i n s n u c l e a r m a t e r i a l i n a d i f f u s e s t a t e , and a f e r t i l i z a t i o n canal i s present.  Figures  Figure  33c-d, Two e x a m p l e s o f s p e r m a t i a c o n t a i n i n g c o n d e n s e d n u c l e a r m a t e r i a l i n c l o s e a s s o c i a t i o n with the f e r t i l i z a t i o n canal. 33e.  a  Figure  33j.  Carpogonium with l a r g e , suspected f u s i o n n u c l e u s ( n o t e empty s p e r m a t i u m and f e r t i l i z a t i o n c a n a l ) .  Figure  33k.  The f i r s t d i v i s i o n o f t h e f u s i o n n u c l e u s c y t o k i n e s i s (see a l s o F i g u r e 3 0 a ) .  Figure  331.  Two the  Figures  Figures  Scale  C a r p o g o n i u m w i t h a t t a c h e d empty s p e r m a t i u m and d e n s e l y s t a i n i n g mass o f s p e r m a t i a l n u c l e a r m a t e r i a l i n the carpogonium c l o s e to the f e r t i l i z a t i o n canal. 3 3 f - i . . S e v e r a l e x a m p l e s of c a r p o g o n i a w i t h empty s p e r m a t i a on t h e i r o u t e r w a l l s and from 1-5 d e n s e l y s t a i n i n g masses o f s p e r m a t i a l n u c l e a r material inside.  prior  to  c a r p o s p o r e s p r o d u c e d by t h e f i r s t d i v i s i o n c a r p o s p o r a n g i u m (see a l s o F i g u r e 30b).  of  33m-o. S u b s e q u e n t s t a g e s o f d i v i s i o n t o p r o d u c e ** ( F i g u r e 33m) o r 8 ( F i g u r e 33o) carpospores.  bar  =  10 um  on  a l l Figures.  200  201  Figure  34.  Diagram s u m m a r i z i n g t h e known m o r p h o l o g i c a l and c y t o l o g i c a l l i f e h i s t o r y of Porphyra g a r d n e r i . I t i s suspected that meiosis occurs i n the conchosporangium a t the time of c o n c h o s p o r e formation.  202  PORPHYRA GARDNERI L I F E  HISTORY  CARPOSPORANGIUM  203  Figure  35.  Porphyra n e r e o c y s t i s spermatium i n a spermatangium near the r e l e a s i n g margin. The s p e r m a t i u m c o n t a i n s abundant s m a l l and l a r g e f i b r o u s v e s i c l e s , a reduced c h l o r o p l a s t , m i t o c h o n d r i a , and c e n t r a l n u c l e u s . ,  204  205  Porphyra n e r e o c y s t i s F i g u r e s 36a-b. T r a n s v e r s e s e c t i o n c f the f o l i o s e t h a l l u s t h r o u g h c a r p o g o n i a and s u s p e c t e d s p e r m a t i a forming f e r t i l i z a t i o n c a n a l s . Figure  36c.  E l e c t r o n micrograph through a suspected canal. Porphyra  of a t r a n s v e r s e s e c t i o n s p e r m a t i u m and fertilization  thuretii  Figure  36d.  T r a n s v e r s e s e c t i o n of the f o l i o s e t h a l l u s a c a r p o g o n i u m and s u s p e c t e d s p e r m a t i u m .  Figure  36e.  T r a n s v e r s e s e c t i o n o f the f o l i o s e t h a l l u s through a carposporangium t h a t has undergone t h e f i r s t division. A s u s p e c t e d s p e r m a t i u m and f e r t i l i z a t i o n canal are present.  Scale  bar =  10  um  on F i g u r e s 3 6 a - b , 36d-e.  through  206  207  Figure  37a.  Porphyra n e r e o c y s t i s spermatia.  n=3.  Figure  37b.  Porphyra n e r e o c y s t i s vegetative f o l i o s e t h a l l u s , n=3.  c e l l s of the  Figure  37c.  Porphyra t h u r e t i i  Figure  37d,e P o r p h y r a n e r e o c y s t i s c a r p o s p o r e . chromosomes v i s i b l e ) .  Figure  37f,g  spermatium.  n=2. 2n=6.  Porphyra n e r e o c y s t i s ccnchosporangial 2n=6.  S c a l e b a r ~ 5 um on a l l F i g u r e s  (only 5 branch,  208  209  GENERAL SUMMARY  Field,  culture  Porphvra s p e c i e s  1.  The  2.  genus  genus  gardneri  major  of 3  epiphytic  results:  is  has t h e r e f o r e  invalid  and  been t r a n s f e r r e d t o  Porphyra.  Porphyra g a r d n e r i .  occurrence,  cn  the  host  morphology,  P. n e r e o c y s t i s  similar  characterized  and  species, basis  species,  P. t h u r e t i i ,  have of  been  their  vegetative  and  3  clearly seasonal  reproductive  growth i n c u l t u r e , and chromosome  numbers.  The s p e r m a t a n g i u m and c a r p o g o n i u m o f  these  3  species  have  division  sequence  been  followed  and  characterized,  in detail.  concluded  that  From  there  and t h e i r these  observations  a r e two b a s i c  types of  I  have  carpogcnial  s p e r m a t a n g i a l m o r p h o l o g y i n t h e genus P o r p h y r a . and  that  the  classical  reported  by  previously  4.  studies  Porphyrella  superficially  3.  cytological  have y i e l d e d f o u r  Porphyrella the  and  The  (1902)  may  of  sexual  not  be  reproduction  has been u n e q u i v o c a l l y  time.  division* as  s e q u e n c e as common  as  thouqht.  occurrence  Porphyra first  Hus  * cruciate  i n t h e qenus  demonstrated  for  the  210  Ss raised  is  the  mere q u e s t i o n s  requiring  1.  always  further  The  than  nature  determines  Does  the  studies  host  light  electron  and  are  of  in  to  the  conchosporangium?  microscope  is  c o n d i t i o n s and for  at  needed  the to  occurs.  physiological  spermatangium  the  Gracilaria  (van  der Meer,  may  provide  1977)  sexuality  spermatangia  and  established.  Gf  Porphyra  Kumford, and  and  and  how  lines  the  carposporangia  of t h o s e  have  1977;  van  der  insight  into  what  various on  particular interest katadai  which  patterns  of  thallus  are  the here  are  species  (Kurogi,  197 2 ) ,  P.  brumalis-  s e v e r a l other species  listed  by  Mumford  which  t h a l l u s and  both  level  along  Todd,  determines  what  induction?  studies  and  elucidate  conchosporogenesis  responsible  been done on  (1975),  needed  t h a t t h i s i s where m e i o s i s  carpogonium  like  topics  relationship.  1  What a r e t h e e n v i r o n m e n t a l  fleer  Major  epiphyte- host'  occur  study  Genetic  answered.  s t u d i e s have  specificity.  Cytological  mechanisms  4.  r e s e a r c h , my  t h e y have  of  meiosis  establish  3.  with  investigation include;  Physiological  2.  case  have t h e s p e r m a t a n g i a  the c a r p o s p o r a n g i a  on t h e  on  one  other.  half  cf  the  211  Can  Pqrphyra  reported well  h y b r i d s be  by  Sutc  (1963),  Such  widespread  of  the  but h i s e x p e r i m e n t s  langia  Bangiophycidae?  (1927),  1978)  with  Based  observations  fuscopurpurea.  Dangeard  and  carpogonia,  Cole  and  and  on  of  McDonald's  the  a  be  (1972) has  the spermatia the  Yabu's diploid  i n Smithora.  fuscopurpurea gametes.  and  HcBride  spermatium  of  of  may  (1967)  the  therefore  gardneri be  thallus  of  the  life  members o f t h e  histories  seem  No  i n the  fertilization  evidence a  of these  Bangiophycidae  algae  would be  of  similarities and  Eangia  functional  boryana  over a c a r p o s p o r e - l i k e c e l l  i n view o f t h i s  of  male  the attachment  Berthold  observed.  report  ultrastructure  reported  alga.  spermatia  certainly  They have some  Erythrotrichia  this  (1909),  In t h e E r y t h r o p e l t i d a c e a e ,  Porphyra  also  of  chromosome number i n t h e  r e p o r t e d on  spermatia  spermatia  canals connecting  o c c u r r i n g i n Bangja.  McBride  (1972)  (personal communication, 3  c a r p o s p o r e s , s e x u a l r e p r o d u c t i o n would  of  not  members  the r e p o r t s of Bosenvinge  of f e r t i l i z a t i o n  fertilization  with  were  were  i s sexual reproduction i n other  ultrastructural  to  results  documented.  How  May,  produced?  of a  (Montagne) filamentous canal  was  re-investigation as w e l l as worthwhile.  other  212  LITEEATUEE CITED  A b b o t t , I . A., and G. J . H o l l e n t e r g . 1976. M a r i n e A l g a e o f C a l i f o r n i a . Stanford U n i v e r s i t y Press, Stanford. xii*827 Pp. A b b o t t , I . A., and ». J , N o r t h . 1972. .Temperature i n f l u e n c e s on f l o r a l c o m p o s i t i o n i n C a l i f o r n i a c o a s t a l w a t e r s , pp. 72-79 i n : K. N i s i z a w a , ed., P r o c . 7 t h I n t . Seaweed Symp. , S a p p o r o , J a p a n 1971. U n i v . Tokyo P r e s s , T o k y o . A n d e r s o n , C. L. 1891. L i s t o f C a l i f o r n i a n o t e s . Zoe 2: 217-225.  marine a l g a e ,  with  i  A r a s a k i , S. 1947. S t u d i e s on t h e w a s t i n g d i s e a s e o f t h e c u l t u r e d l a v e r s (Pcrphyra tenera) [ i n Japanese}. B u l l . J a p . Soc. , S c i . F i s h . 13: 7 4-90. A r a s a k i , S. 1960. A c h y t r i d e a n p a r a s i t e on t h e P o r p h y r a f i n Japanese, E n g l i s h a b s t r a c t ] . B u l l . Jap. Sec. S c i . F i s h . 26: 543-548. A r a s a k i , S.,1962. S t u d i e s on t h e a r t i f i c i a l t e n e r a K j e l l m . I I I . On t h e r e d w a s t i n g P o r p h y r a . e s p e c i a l l y on t h e p h y s i o l o g y fungus Pythium s p . nov. [ i n Japanese, Noden K e n k y u j o Ho [ J . A g r . L a b . A b i k o ,  c u l t u r e o f Porphyra disease of of the causal English abstract]. J a p a n ] 3: 87-93.  A r a s a k i , S. 1974. A c o m p a r i s o n i n t h e p h e n o l o g y o f t h e l i t t o r a l a l g a , P p r p h y r a . on t h e c o a s t o f J a p a n and c n t h e west c o a s t o f t h e U.S.A. p. A59 i n : A b s t r a c t s o f P a p e r s , 8 t h Int. Seaweed Symposium, Bangor, H a l e s . A r a s a k i , S., A. I n o u y e and Y. K o c h i . 1960. The d i s e a s e o f t h e c u l t u r e d Pprphyra. with s p e c i a l r e f e r e n c e to the c a n c e r - d i s e a s e a n d t h e c h y t r i d - d i s e a s e which o c c u r r e d a t t h e c u l t u r e f i e l d i n Tokyo-Bay d u r i n g 1959-1960 f i n Japanese, E n g l i s h a b s t r a c t ] . B u l l . J a p , Soc. S c i . F i s h . 26: 1074-1081. B a t t e r s , E. A. L. 1892. On C o n c h o c e l i s . a new genus o f p e r f o r a t i n g a l g a e , pp. 25-28; p i , 8. i n : G. M u r r a y , P h y c o l o g i c a l M e m o i r s , p a r t 1. D u l a n and C o . , London. B e r t h o l d , G. 1882. D i e B a n g i a c e e n d e s g o l f e s von N e a p e l und d e r Angrenzenden M e e r e s - A b s c h n i t t e . Fauna und F l o r a d e s G o l f e s von N e a p e l , e t c . 8: 1-28; f i g . 1. Bird,  C. J . 1973. A s p e c t s o f t h e l i f e h i s t o r y and e c o l o g y o f P o r p h y r a l i n e a r i s ( B a n g i a l e s , fihodophyceae) i n n a t u r e . Can. J . B o t . 51: 2371-2379.  B i r d , C. J . , L. C.-M. Chen and J . M c L a c h l a n . 19 72. The c u l t u r e c f P o r p h y r a l i n e a r i s ( B a n g i a l e s , R h o d o p h y c e a e ) . Can. J .  213  Bot.  50:  1859-1863.  B l a n k i n s h i p , J . W., h i s t o r y of the  and C. A. K e e l e r . 189 2. On t h e n a t u r a l F a r a l l o n I s l a n d s . Zoe 3: 144-165.  B o u r n e , V. L., E. Conway and K. C o l e . 1970. On t h e u l t r a s t r u c t u r e of p i t c o n n e c t i o n s i n t h e c o n c h o c e l i s o f t h e r e d a l g a P o r p h y r a p e r f o r a t a 3. Ag. P h y c o l o g i a 79-81. Ercwn, D. L. 1969. U l t r a s t r u c t u r e o f t h e r e d a l g a B a t r a c h o s p e r m u m . Ph.D. t h e s i s . U n i v e r s i t y of D a v i s , vi+154 pp.  phase 9:  California,  E r y a n , J . H. D., and B. R. B r i n k l e y . 1963. An e l e c t r o n m i c r o s c o p e s t u d y of t h e s i l v e r - S c h i f f r e a c t i o n . J . B i o l . 19: 10A ( A b s t r . ) .  Cell.  B r y a n , J . H. D., and B. R. B r i n k l e y . 1964. A s i l v e r - a l d e h y d e r e a c t i o n f o r s t u d i e s o f chromosome u l t r a s t r u c t u r e . Q. J . H i c r o s c . S c i . 105: 367-374. Chemin, E. 1927. L e s mouvetnents a m i b o i d e s des s p o r e s c h e z g u e l g u e s F l o r i d e e s . C. R. Soc, B i o l . P a r i s 97: 1677-1679. Chemin, E. 1929. L e s mouvements a m i b o i d e s des s p o r e s de Bonnemaisonia a s p a r a g o i d e s A g . Am. J . P h y s i o l . 90:  312.  Chen, L . C-M., I . E d e l s t e i n , E.,Ogata and J . M c L a c h l a n . , 1 S 7 0 . The l i f e h i s t o r y o f P o r p h y r a m i n i a t a . C a n . J . B o t . 48: 385-389. C h i h a r a , fl. 1975. R h o d o p h y t a , t h e i r l i f e - h i s t o r i e s , pp.,137-150 i n : J . T o k i d a and H, H i r o y u k i , e d s . . A d v a n c e o f P h y c c l c g y i n J a p a n . Junk, The Hague. Cole,  K. 1S72a. O b s e r v a t i o n s on the l i f e h i s t o r y o f B a n g i a f u s c o p u r p u r e a . flem. Soc. B e t . F r . 1972: 231-236.  C o l e , K. 1972b. Some e l e c t r o n m i c r o s c o p i c o b s e r v a t i o n s on t h e c u l t u r e d c o n c h o c e l i s phase o f P o r p h y r a s p e c i e s , pp. 157-166 i n : I . A. A b b o t t and M. K u r o g i , e d s . , C o n t r i b u t i o n s t o the S y s t e m a t i c s o f B e n f h i c Marine Algae of t h e N o r t h P a c i f i c . J a p . Soc. P h y c o l . , Kobe, J a p a n . Cole,  K., and E. Conway, 1975. P h e n e t i c i m p l i c a t i o n s of s t r u c t u r a l f e a t u r e s of the p e r e n n a t i n g phase i n the h i s t o r y o f P o r p h y r a and Batngia ( B a n g i o p h y c e a e , R h o d o p h y t a ) . P h y c o l o g i a 14: 239-245.  life  C o l l i n s , F. S. 1913. The m a r i n e a l g a e o f V a n c o u v e r I s l a n d . Canada G e o l . S u r v e y , V i c t o r i a M e m o r i a l Museum B u l l . Nc. 13: 99-137.  1,  C o l l i n s , F. S., I . Hoiden and a . A. S e t c h e l l . 1919. P h y c o t h e c a B o r e a l i - A m e r i c a n a , F a s c i c l e 46, No. 2297. M a i d e n , Mass.  214  Conway, E. 1964a. A u t e c o l o g i c a l s t u d i e s o f t h e genus P o r p h y r a : I . The s p e c i e s f o u n d i n B r i t a i n . B r i t . P h y c o l . B u l l . 2: 342-348. Conway, E. 1964b. A u t e c o l o g i c a l s t u d i e s o f t h e genus P o r p h y r a : I I . P o r p h y r a u m b i l i c a l i s ( I . ) J . Ag. B r . P h y c o l . B u l l . 2: 349-363. ~ Conway, E . 1964c. C o n t r i b u t i o n s t o o u r knowledge o f t h e l i f e - h i s t o r y o f P o r p h y r a s p e c i e s , pp. 66-69 f d i s c u s s i o n p. 70 ] i n : D. de V i r v i l l e and J . Feldmann, e d s . , P r o c . 4 t h I n t . Seaweed Symp., B i a r r i t z 1961, Pergamon P r e s s , M a c M i l l a n C o . , New York. Conway, E. 1966. J u v e n i l e s t a g e s i n t h e genus P o r p h y r a . pp. 101-105 i n : E. G. Young and J . L . M c L a c h l a n , e d s . , P r o c . 5 f h I n t . Seaweed Symp., H a l i f a x 1965. Pergamon P r e s s , Oxford, Toronto, e t c . Conway, E. 1973. P o r p h y r a (Afcstr.).  i n the P a c i f i c .  Br. Phycol.  J . 8: 207  Conway, E . , and K. C o l e . 1973. O b s e r v a t i o n s on an u n u s u a l form o f r e p r o d u c t i o n i n P o r p h y r a (Rhodophyceae, B a n g i a l e s ) . P h y c o l o g i a 12: 213-225. Conway, E . , and K. C o l e . 1977. S t u d i e s i n t h e B a n g i a c e a e : s t r u c t u r e and r e p r o d u c t i o n o f t h e c o n c h o c e l i s o f P o r c h v r a and Bang,ia i n c u l t u r e { B a n g i a l e s , R h o d o p h y c e a e ) . P h y c o l o g i a 16: 205-216. Conway, E . , and K. C o l e . I n p r e s s . Some c y t o l o g i c a l c o n s i d e r a t i o n s o f l i f e - h i s t o r i e s i n t h e genus P o r p h y r a . i n : G. F o g g , e d . , P r o c , 8 t h I n t . Seaweed Bangor 1974?  Symp.,  Conway, E., and A. P. W y l i e . 1972. S p o r e o r g a n i z a t i o n and r e p r o d u c t i v e modes i n two s p e c i e s o f P o r p h y r a f r o m New Z e a l a n d , pp. 105-107 i n : K. N i s i z a w a , e d . , P r o c , 7 t h I n t . Seaweed Symp., S a p p o r o J a p a n . 1971. Univ. T o k y o P r e s s , Tokyo. Conway, E . , T. F. Mumford J r . , and R. F. S c a g e l . 1975. The g e n u s P o r p h y r a i n E r i t i s h C o l u m b i a and W a s h i n g t o n . S y e s i s 8: 185-244 [ I s s u e d A p r i l , 1976J. C r a i g i e , J . S., a n d J . M c L a c h l a n . , 1 9 6 4 . E x c r e t i o n o f c o l o u r e d u l t r a v i o l e t - a b s o r b i n g s u b s t a n c e s by marine a l g a e . C a n . J . B o t . 42: 23-33. D a n g e a r d , P. 1927. H e c h e r c h e s s u r l e s B a n g i a e t l e s P o r p h y r a . B o t a n i s t e 18: 183-244, 12 p l . D a n g e a r d , P. 1929. S u r l e s c h r o m a t o p h o r e s de 1 * A c r o c h a e t i u m c a e s j i t o s u m N a e g e l i . B u l l . S o c . B o t . F r . 76: 1043-1048.  215  Dawes, C. J . 1971. B i o l o g i c a l T e c h n i g u e s i n E l e c t r o n M i c r o s c o p y , B a r n e s and N o b l e , I n c . , H a r p e r and Row, Y o r k , x i v * 1 9 3 pp.  New  Dawson, E. Y. 1944. The m a r i n e a l g a e o f t h e G u l f o f C a l i f o r n i a . A l l a n Hancock P a c i f i c E x p e d . 3: 189-453. Dawson, E. Y. 1951. A f u r t h e r vegetation along P a c i f i c Res. 10: 39-58.  s t u d y o f u p e l l i n g and a s s o c i a t e d B a j a C a l i f o r n i a , M e x i c o . J . Mar.  Dawson, E. Y. 1952. M a r i n e r e d a l g a e o f P a c i f i c M e x i c o , P a r t I . B a n g i a l e s t o C o r a l l i n a c e a e Subf. C o r a l l i n c i d e a e . A l l a n Hancock P a c i f i c Exped. 17: 1-239, [ I s s u e d Feb. 1953]. Dawson, E. Y. 1961. A g u i d e t o t h e l i t e r a t u r e and d i s t r i b u t i o n s of P a c i f i c b e n t h i c algae frcm Alaska t o t h e Galapagos I s l a n d s . Pac. S c i . 15: 370-461. D e r b e s , A., a n d A . - J . - J . S c l i e r . 1856. Memoire s u r g u e l g e s p o i n t s de l a p h y s i o l o g i e des a l g u e s . Ccmp. Rend. Acad. S c i . P a r i s , s u p p l . 1: 1-120; p i s . 1-23. Dixon,, P. S. 1970. The R h o d o p h y t a : Some a s p e c t s o f t h e i r b i o l o g y . I I . O c e a n o g r . Mar. B i o l . Ann. Rev. 8: 307-352. D i x o n , P. S. 1973. B i o l o g y o f R h o d o p h y t a . O l i v e r E d i n b u r g h , x i i i + 2 8 5 pp.  and Boyd,  D i x c n , P. S., and W. N. R i c h a r d s o n . 1969. The l i f e h i s t o r i e s o f B a n g i a and P o r p h y r a and t h e p h o t o p e r i o d i c c o n t r o l c f s p o r e p r o d u c t i o n , pp. 133-139 i n : R, S a r g a l e f , e d . , P r o c . 6 t h I n t . Seaweed Symp., S a n t i a g o de C o m p o s t e l a 1968. S u b s e c r e t a r i a de l a m a r i n a m e r c a n t e , M a d r i d . Doty,  M. S..1947. M a r i n e a l g a e o f O r e g o n . P a r t F a r l o w i a 3: 159-215.  I I . Rhodophyta.  Drew, K. M. 1949. C o n c h o c e l i s - p h a s e i n t h e l i f e - h i s t c r y c f P o r p h y r a u m b i l i c a l i s (L.) Ku'tz. N a t u r e (London) 164: 748-749. Drew, K. M. 1951. R h o d o p h y t a . pp. 167-191 i n : G, M. S m i t h , e d . , Manual o f P h y c o l o g y . C h r o n i c a B o t a n i c a Co. , JJaltham, Mass. Drew, K. M. 1954a. S t u d i e s i n t h e B a n g i o i d e a e . I I I . The l i f e - h i s t o r y o f P o r p h y r a u m b i l i c a l i s (L.) K'utz. v a r . l a c i n i a t a ( L i g h t f . ) J . Ag. Ann. B o t . (London) N. S. 18: 183-211; p i s . 9-12. Drew, K.M. 1954b. L i f e - h i s t o r y 173: 1243-1244.  of P o r p h y r a . Nature  (London)  Drew, K. M. 1956. R e p r o d u c t i o n i n t h e B a n g i o p h y c i d a e . 22: 553-611.  B o t . Rev.  216  Drew, K...8., a n d K. S. B i c h a r d s . 1953, S t u d i e s i n t h e Bangioideae. I . The conchocelis-phase of Porphvra s c . i n P c l l i c i p e s c o r n u c o p i a Leach a t R o s c o f f . J . L i n n . S o c . London, B o t . 60: 84-87; p i . 1. D r i n g , M . J . 1967. E f f e c t s o f d a y l e n g t h on g r o w t h and reproduction o f t h e conchocelis-phase c f Porphyra J . Mar. B i o l , a s s o c . 0. K. 47: 501-510.  tenera.  E v a n s , L . V., G. P. B o l w e l l , J . a. C a l l o w and M. E. C a l l o w . 1977. Egg-sperm r e c o g n i t i o n i n f u c o i d s . J . P h y c c l . S u p p l . t o 13: 80 ( a b s t r . ) . F e d e r , K., and T. P. O ' B r i e n . 1968. P l a n t m i c r o t e c h n i q u e : Seme p r i n c i p l e s a n d new methods, am, J . B o t . 55: 123-142. F e t t e r , R. 1977. Bed a l g a l r e p r o d u c t i o n v i a s p e r m a t i a c o n t a i n i n g s l i m e s t r a n d s , J . P h y c o l . S u p p l . t o 13: 80 (abstr.). F r i t s c h , F. E. 1945. T h e S t r u c t u r e and R e p r o d u c t i o n a l g a e . V. 2. C a m b r i d g e U n i v . P r e s s , Cambridge, 1 map.  of the xiv+939 pp,  F u k u h a r a , E . 1968, S t u d i e s on t h e taxonomy and e c o l o g y o f P o r p h y r a o f H o k k a i d o and i t s a d j a c e n t waters f i n J a p a n e s e , E n g l i s h summary]. B u l l , H o k k a i d o Reg. F i s h . Res. L a b . 34:  40-99.  F u l l e r , M, S., B. L e w i s and P. Cook. 1966. O c c u r r e n c e o f P y t h i u j i s p . on t h e m a r i n e a l g a P o r p h y r a . M y c o l o g i a 58:  313-3T8.  G i r a u d , A., and F. Magne. 1968. L a p l a c e de l a meiose d a n s l e c y c l e de d e v e l c p p e m e n t d e P o r p h y r a u m b i l i c a l i s . C. R. a c a d . S c i . P a r i s , S e r . D, 267: 586-588. G o f f , L. J . 197 5. A d e v e l o p m e n t a l , p h y s i o l o g i c a l and s t r u c t u r a l study o f t h e rhodophycean a l l o p a r a s i t e H a r v e v e l l a m i r a b i l i s ( C h o r e o c o l a c e a e , C r y p t o n e r a i a l e s ) . Ph.B. t h e s i s , U n i v e r s i t y o f B r i t i s h C o l u m b i a , V a n c o u v e r . 239 pp. G r a v e s , J . M. 1955. L i f e - c y c l e o f P o r p h y r a N a t u r e (London) 175: 393-394.  capensis  Kutz.  G r a v e s , J . M. 1969. T h e genus P o r p h y r a o n S o u t h A f r i c a n c o a s t s : I. Observations on t h e autecology o f Porphyra c a p e n s i s s e n s u I s a a c (1957), i n c l u d i n g a d e s c r i p t i o n o f dwarf p l a n t s . J . S. a f r . B o t . 35: 343-362. Grubfc, V. M. 1923. The a t t a c h m e n t s o f P o r p h y r a jam b i I l e a l i s , (1.) J . ag. ann. B o t . (London) N. S. 37: 131-140; p i  1.  Grubfc, V. M, 1924, O b s e r v a t i o n s o n t h e e c o l o g y and r e p r o d u c t i o n °f P o r p h y r a u m b i l i c a l i s (L.) J , ag. Rev. A l g o l . 1:  223-234.  217  Grubb, V. M. 1925. The male o r g a n s o f t h e F l o r i d e a e . J . L i n n . Soc. London, B o t . 47: 177-255. Gunawardena, P., and F. W i l l i a m s o n . 1974. S t r u c t u r e and c o m p o s i t i o n o f t h e c e l l w a l l s , i n t e r c e l l u l a r m a t r i x and o u t e r c u t i c l e of P o r p h y r a u m b i l i c a l i s . pp. B7 i n : a b s t r a c t s o f p a p e r s , 8 t h I n t . Seaweed Symp., Bangor, Wales. Hamel, G. 1924. F l o r i d e e s de F r a n c e , 1-46 [ R e p r i n t page n u m b e r s ] .  Bangiales.  Rev.  Algol.  1:  H a n i c , L . A., and J . S . C r a i g i e 1969. S t u d i e s on t h e a l g a l c u t i c l e . J . P h y c o l . 5: 89-102. H a r l i n , M. , M. 1S73a. ' O b l i g a t e * a l g a l e p i p h y t e : S m i t h o r a naiadum grows on a s y n t h e t i c s u b s t r a t e . J . P h y c o l . 9: 230-232. H a r l i n , M. M. 1973b. T r a n s f e r o f p r o d u c t s between e p i p h y t i c marine a l g a e and h o s t p l a n t s . J . P h y c o l . 9: 243-248. Hawkes, M. W. 1977a, A f i e l d , c u l t u r e and c y t o l o g i c a l s t u d y o f P o r p h y r e l l a g a r d n e r i Smith 6 H o l l e n b e r g ( B a n g i a l e s ) . J . P h y c o l s u p p l . t o 13: 28 ( A b s t r . ) . Hawkes, M. W. 1977b. A f i e l d , c u l t u r e and c y t o l o g i c a l s t u d y o f P o r p h y r a g a r d n e r i (Smith 6 H o l l e n b e r g ) comb. nov., ( = P o r p h y r e l l a g a r d n e r i Smith & H o l l e n b e r g ) , ( B a n g i a l e s , R h o d o p h y t a ) . P h y c o l o g i a 16: 457-469. Hawkes, M. W. 1978. S e x u a l r e p r o d u c t i o n i n P o r p h y r a g a r d n e r i (Smith e t H o l l e n b e r g ) Hawkes ( B a n g i a l e s , B h o d o p h y t a ) . P h y c o l o g i a 17: i n p r e s s . H e n r y , E. C. 1977. A method f o r o b t a i n i n g r i b b o n s o f s e r i a l s e c t i o n s o f p l a s t i c embedded s p e c i m e n s . S t a i n T e c h . 52: 59-60. Hoek, C. van den. 1958. The a l g a l m i c r o v e g e t a t i o n i n and cn b a r n a c l e - s h e l l s c o l l e c t e d a l o n g t h e butch and F r e n c h c o a s t s . Blumea 9: 206-214. H o l l e n b e r g , G . J . 1945. New m a r i n e a l g a e from s o u t h e r n C a l i f o r n i a . I I I . Am. J . B o t . 32: 447-451. H o l l e n b e r g , G . J . 1958. C u l t u r e s t u d i e s o f marine a l g a e . I I I . P o r p h y r a p e r f o r a t a . Am. J . B o t . 45: 653-656. H o l l i s t e r , H. J . 1968. Sea s u r f a c e t e m p e r a t u r e and s a l i n i t y a t shore s t a t i o n s along t h e B r i t i s h Columbia c o a s t d u r i n g 1966 [ u n p u b l i s h e d MS]. F i s h . R e s . B o a r d Can. T e c h . R e p t . No: 82: i i * 3 9 pp. Howe, M. A. 1893. A month E r y t h e a 1: 63-68.  on t h e s h o r e s  of aonterey  Bay.  218  Hus, H . T . A. 1900. P r e l i m i n a r y n o t e s on w e s t - c o a s t Zoe 5: 61-70. Bus,  Porphyras.  H. T. A. 1902. An a c c o u n t o f t h e s p e c i e s o f P o r p h y r a f o u n d on t h e P a c i f i c c o a s t o f N o r t h A m e r i c a . P r o c . C a l i f . Acad. S c i . 2: 173-240; p i s . 20-22.  Imada, 0., Y. S a i t o and K - I . T e r a m o t o . 1972. A r t i f i c i a l c u l t u r e o f l a v e r . pp. 358-363 i n : K. N i s i z a w a , e d . , P r o c . ,7th I n t . Seaweed Symp. , S a p p o r o , J a p a n 1971. U n i v . Tokyo P r e s s , Tokyo. I s h i k a w a , H. 1921. C y t o l o g i c a l s t u d i e s on P o r p h v r a K j e l l m . B o t . Mag. Tokyo 35: 206-218; p i . 4.  tenera  I w a s a k i , H. 1961. The l i f e - c y l e o f P o r p h y r a t e n e r a i n B i o l . B u l l . Hoods H o l e , Mass. 121; 173-187."  vitro.  I w a s a k i , H. 1965. S t u d i e s on t h e p h y s i o l o g y and e c o l o g y o f P o r p h y r a t e n e r a [ i n J a p a n e s e , E n g l i s h summary]. J . F a c . F i s h . Anim. Husb. B i r o s h i m a U n i v . 6: 133-211. I w a s a k i , H. 1967. N u t r i t i o n a l s t u d i e s o f t h e e d i b l e seaweed Porphyra tenera. I I . N u t r i t i o n of C o n c h o c e l i s . J . Phycol. 3: 30-34. I w a s a k i , H., and C. M a t s u d a i r a . 1958. C u l t u r e o f a l a v e r , P o r p h y r a t e n e r a K j e l l m . I . P r e l i m i n a r y r e s e a r c h on c u l t u r a l c o n d i t i o n s [ i n J a p a n e s e , E n g l i s h summary]. B u l l . J a p . S o c . S c i . F i s h . ~ 24: 396-401 . I w a s a k i , H.^ and C. M a t s u d a i r a . 1963. O b s e r v a t i o n on t h e e c o l o g y and r e p r o d u c t i o n c f f r e e - l i v i n g c o n c h o c e l i s c f P o r p h y r a t e n e r a . B i o l . B u l l , woods H o l e , Mass. 124: 268-276. I w a s a k i , H.» and N. S a s a k i . 1972. The C o n c h o c e l i s - p h a s e o f P p r p h y r a s u b o r b i c u l a t a f o r m a l a t i f p i j a . p p . 36 4-367 in,: N i s i z a w a , e d . , P r o c . 7 t h I n t . Seaweed Symp., S a p p o r o , J a p a n 1971. U n i v . T o k y o P r e s s , T o k y o . . . . .  K.  /  J a n c z e w s k i , E. 1873. E t u d e s a n a t o m i g u e s u r l e s P o r p h y r a • e t s u r l e s p r o p a g u l e s du S p h a c e l a r l a c i r r o s a . Ann. S c i . Nat. s e r . 5, (Bot.) 17: 241-260; p i . 19. J o f f e ' , B. 1896. O b s e r v a t i o n s s u r l a f e c o n d a t i o n d e s B u l l . S e c . B o t . F r a n c e 43; 143-146; p i . 2.  Bangiacees.  J o h a n s e n , fl..W. 1971. E f f e c t s o f e l e v a t i o n c h a n g e s on b e n t h i c a l g a e i n P r i n c e W i l l i a m Sound, pp. 35-68 i n : The G r e a t A l a s k a E a r t h g u a k e o f 1964 - B i o l o g y . Nat. Acad, c f S c i e n c e s , U.S.A., W a s h i n g t o n . J u r a n d , A., K. D e u t s c h and A. E . G . Dunn.,1958. A p p l i c a t i o n o f t h e s i l v e r - F e u l g e n r e a c t i o n i n e l e c t r o n m i c r o s c o p y . J . R. M i c r o s c . Soc. 78: 46-52; p i s . 6-7.  219  Kazama, I . , and M, S. F u l l e r . 1970. U l t r a s t r u c t u r e o f P o r c h v r a p e r f o r a t a i n f e c t e d w i t h P y t h i u m marinum, a m a r i n e f u n g u s . Can. J . B o t . 48: 21C3-2107. K i r i t a , a . 1970. The e f f e c t o f Germanium d i o x i d e , a d i a t o m - e l i m i n a t i n g c h e m i c a l , on t h e growth o f f r e e - l i v i n g C o n c h o c e l i s of P o r p h y r a y e z o e n s i s f i n J a p a n e s e , E n g l i s h summary]. B u l l . J a p . S o c . P h y c o l . 18: 167-170. Kito,  H. 1966. C y t o l o g i c a l s t u d i e s o f s e v e r a l s p e c i e s o f P o r p h y r a . I . M o r p h o l o g i c a l and c y t o l o g i c a l o b s e r v a t i o n s on a s p e c i e s o f P o r p h y r a e p i p h y t i c on G r a t e 1 p u p i a filicina v a r . p o r r a c e a ( M e r t . ) Howe [ i n J a p a n e s e , E n g l i s h summary]. B u l l . F a c . F i s h . , H o k k a i d o U n i v . 16: 206-208; p i s . 1-5.  K i t e , H. 1967. C y t o l o g i c a l s t u d i e s o f s e v e r a l s p e c i e s o f Porphyra. I I . M i t o s i s i n carpospore-germlings of Porphyra y e z o e n s i s [ i n J a p a n e s e , E n g l i s h summary]. B u l l . Fac. F i s h . , H o k k a i d o U n i v . 18: 201-202; p i s . 1-2. K i t e , H. 1968. C y t o l o g i c a l s t u d i e s o f s e v e r a l s p e c i e s o f P o r p h y r a . I I I . Chromosome number o f P p r p h y r a t e n e r a K j e l l m a n [ i n J a p a n e s e , E n g l i s h summary]. B u l l . Tohoku F i s h . Ees. L a b . 28: 137-140; p l . 1. K i t o , H. 1974. C y t o l o g i c a l o b s e r v a t i o n s on the conchocelis-phase i n three s p e c i e s of Porphyra f i n J a p a n e s e , E n g l i s h summary]. B u l l . Tohoku Beg. F i s h . L a b . 33: 101-117; p i s . 1-7. Kito,  H,, E. O g a t a and J . M c L a c h l a n . 1971. C y t c l o g i c a l o b s e r v a t i o n s on t h r e e s p e c i e s o f P o r p h y r a f r o m t h e A t l a n t i c . /Bot. Mag. Tokyo 84; 141-148; p i s . 1-4.  Kite,  H., H. Yabu and J . T o k i d a . 1967. The number o f chromosomes i n seme s p e c i e s c f P o r p h y r a . B u l l . Fac. H o k k a i d o U n i v . 18: 59-60; p i s . 1-2.  K l e i n , B. M., and D. T, K l e i n . 1S7C. Science. Natural History Press, York, xi+756 pp.  Beg.  fies.  Fish..  R e s e a r c h Methods i n P l a n t Garden C i t y , Sew  Knaggs, F. w. 1966a. S o l e o f l i g h t e n e r g y i n t h e i n i t i a t i o n and d e v e l o p m e n t o f t e t r a s p o r a n g i a on c u l t u r e d s p e c i m e n s of a r e d a l g a l s p e c i e s . N a t u r e (London) 212: 431. Knaggs, F. W. 1966b. B h o d o c h o r t o n purpureum ( L i q h t f . ) B o s e n v i n g e . O b s e r v a t i o n s on t h e r e l a t i o n s h i p between r e p r o d u c t i c n and e n v i r o n m e n t . I , The r e l a t i o n s h i p between t h e e n e r g y o f i n c i d e n t l i g h t and tetrasporangium p r o d u c t i o n . Nova H e d w i g i a Z. K r y p t o g a m e n k d . 11: 405-411. Knaggs* F. H. 1967. R h p d c c h p r t o n f l o r i d u l u m ( D i l l w n . ) Nag. ( O b s e r v a t i o n s on t h e r e l a t i o n s h i p between r e p r o d u c t i c n e n v i r o n m e n t . I ) . Nova H e d w i g i a Z. Kryptogamendk. 14: 31-38; p i s . 32-38.  and  220  Kornmann, P. 1960. Vcn C o n c h o c e l i s zu P o r p h y r a . H e l g o l . M e e r e s u n t e r s , 7: 189-193.  Wiss.  Kcrnmann, P. 1961a. D i e e n t w i c k l u n g von P o r p h y r a l e u c c s t i c t a im k u l t e r v e r s u c h . H e l g o l . H i s s . M e e r e s u n t e r s . 8: 167-175. Kornmann, P. 1961b. Z u r k e n n t n i s d e r P o r p h y r a - a r t e n von H e l g o l a n d . H e l g o l . H i s s . M e e r e s u n t e r s . 8: 176-192. K r i s h n a m u r t h y , V. 1959. C y t o l o g i c a l i n v e s t i g a t i o n s o f P o r p h y r a u m b i l i c a l i s <L.) Ku'tz. v a r . l a c i n i a t a ( L i g h t f . ) J . Ag. Ann. B o t . {London), N • S. 23: 117-176; p i s . 1-2. K r i s h n a m u r t h y , V. 1969a. The C o n c h o c e l i s phase o f t h r e e s p e c i e s °f P o r p h y r a i n c u l t u r e . J . P h y c o l . 5: 42-47. K r i s h n a m u r t h y , V. 1969b. On two s p e c i e s o f P o r p h y r a from San J u a n I s l a n d , W a s h i n g t o n , pp. 225-234 i n : B. M a r g a l e f , e d . , P r o c . 6 t h I n t . Seaweed Symp., S a n t i a g o de C o m p o s t e l a 1968. S u b s e c r e t a r i a de l a m a r i n a m e r c a n t e , K r i s h n a m u r t h y , V. 1972. A r e v i s i o n o f t h e s p e c i e s o f t h e a l g a l genus P o r p h y r a o c c u r r i n g on t h e P a c i f i c c o a s t o f N o r t h A m e r i c a . P a c . S c i . 26: 24-49. K r i s h n a m u r t h y , V. 1977. O b s e r v a t i o n s on t h e c y t o l o g y o f some s p e c i e s o f Porphyra from t h e P a c i f i c c o a s t o f North A m e r i c a w i t h r e f e r e n c e t o t h e l i f e - h i s t o r y o f t h e genus. J . P h y c o l . S u p p l . t c 13: 37 ( A b s t r . ) . K u g r e n s , P. 1974. L i g h t and e l e c t r o n m i c r o s c o p i c s t u d i e s cn t h e development and l i b e r a t i o n of J a n c z e w s k i a g a r d n e r i S e t c h . s p e r m a t i a ( B h o d o p h y t a ) . P h y c o l o g i a 13: 295-306. K u g r e n s , P., and J . A. West. 1972. U l t r a s t r u c t u r e c f s p e r m a t i a l development i n the p a r a s i t i c r e d a l g a e L e y r i n q i e l l a g a r d n e r i and E r y t h r o c y s t i s s a c c a t a . J . P h y c o l . 8: 33 1-343. K u g r e n s , P., and J . A. West. 1973. The u l t r a s t r u c t u r e o f carpospore d i f f e r e n t i a t i o n i n the p a r a s i t i c red alga L e v r i n g i e l l a g a r d n e r i ( S e t c h . ) K y l i n . P h y c o l o g i a 12: 16 3-173. K u g r e n s , P., and J . A. West. 1974. The u l t r a s t r u c t u r e o f c a r p o s p o r o g e n e s i s i n the marine h e m i p a r a s i t i c r e d a l g a l£Yth.tocy.stis s a c c a t a . J . P h y c o l . 10: 139-147. K u n i e d a , H. 1939. On t h e l i f e - h i s t o r y o f P o r p h y r a t e ne r a K j e l l m a n . J . C o l l . A g r i c . Imp. U n i v . Tokyo 14: 377-405; p i s . 14-34. K u r o g i , H. 1953a. S t u d y o f t h e l i f e - h i s t o r y c f P o r p h y r a . I . The g e r m i n a t i o n and development of c a r p o s p o r e s [ i n Japanese, E n g l i s h summary]. B u l l . Tohcku Beg. F i s h . Bes. L a b . 3: 67-103; p i s . 1-3.  221  K u r o g i , M. 1953b. On t h e l i b e r a t i o n o f monospores f r o m t h e f i l a m e n t o u s t h a l l u s (ConchoceJ.is- s t a g e ) o f P o r ph v r a t e n e ra K j e l l m . [ i n Japanese, E n g l i s h summary]. B u l l . Tohoku Beg. F i s h . B e s . L a b . 2: 104-108. K u r o g i , M. 1956. L i f e - h i s t o r y c f P o r p h y r a , e s p e c i a l l y on s o - c a l l e d "summer p l a n t l e t s " [ i n J a p a n e s e ] . B u l l . J a p . S o c . P h y c o l . 4: 13-18. K u r o g i , M. 1959. I n f l u e n c e s o f l i g h t on t h e g r o w t h and m a t u r a t i o n o f C o n c h o c e l i s - t h a l l u s o f Porphvra I . E f f e c t of p h o t o p e r i o d on the f o r m a t i o n o f m o n o s p o r a n g i a and l i b e r a t i o n o f monospores (1) [ i n Japanese, E n g l i s h summary]. B u l l . Tohoku Reg. F i s h . B e s . L a b . 15: 33-42. K u r o g i , H . 1961. S p e c i e s o f c u l t i v a t e d P o r p h y r a s and t h e i r l i f e h i s t o r i e s . (Study o f t h e l i f e h i s t o r y c f P o r p h y r a * I I . ) [ i n Japanese, E n g l i s h summary]. B u l l . Tohoku Beg. F i s h . Bes. L a b . , 1 8 : 1-115; p i s . 1-39. K u r o g i , M. 1972. S y s t e m a t i c s o f P o r p h y r a i n Japan, pp. 167-191; pis. 1-4 i n : I . A. A b b o t t and M. K u r o g i , e d s . . C o n t r i b u t i o n s to the S y s t e m a t i c s o f Benthic Marine Algae of t h e N o r t h P a c i f i c . J a p . S o c . P h y c o l , Kobe. K u r c g i , M., and K . A k i y a m a . 1965. I n f l u e n c e s o f l i g h t on t h e g r o w t h and m a t u r a t i o n o f C o n c h o c e l i s - t h a l l u s o f P o r p h y r a . I V . E f f e c t o f l i g h t i n t e n s i t y on t h e monospore l i b e r a t i o n [ i n Japanese, E n g l i s h summary]. B u l l . Tohoku Reg. F i s h . R e s . L a b . 25: 171-177. K u r o g i , M., and K. Akiyama. 1966. E f f e c t s o f w a t e r t e m p e r a t u r e on t h e growth and m a t u r a t i o n o f C o n c h o c e l i s - t h a l l i i n s e v e r a l s p e c i e s o f P o r p h y r a [ i n Japanese, E n g l i s h summary]. B u l l . Tohoku Reg. F i s h . Bes. L a b . 26: 77-89. K u r c g i , M., and K. H i r a n o . 1955a. I n f l u e n c e s o f d e s i c c a t i o n , s a l i n i t y o f t h e w a t e r and i l l u m i n a t i o n on t h e C o n c h o c e l i s - t h a l l u s o f P o r p h y r a E n g l i s h summary 1. B u l l . Tohoku Reg. F i s h . Res. L a b . 4: 262-278. K u r c g i , M., a n d K. H i r a n o . 1955b. On t h e l i b e r a t i o n o f m c o o s p o r e s from t h e f i l a m e n t o u s t h a l l u s ( C o n c h o c e l i s - s t a g e ) o f P o r p h y r a t e n e r a K j e l l m . (2) The d a i l y p e r i o d i c i t y o f t h e l i b e r a t i o n [ i n Japanese, E n g l i s h summary]. B u l l . T o h o k u Reg. F i s h . R e s . L a b . 4: 279-282. K u r o g i , M., and K. H i r a n o . 1956a. On t h e l i b e r a t i o n o f monospores from C o n c h o c e l i s - p h a s e o f P o r p h y r a . ( E x p e r i m e n t i n t h e sea) [ i n Japanese, E n g l i s h summary]. B u l l . Tohoku Reg. F i s h . R e s . L a b . 8: 27-44. K u r o g i , M., and K. H i r a n o . 1956b. I n f l u e n c e s o f w a t e r t e m p e r a t u r e cn t h e g r o w t h , f o r m a t i o n o f m o n o s p o r a n g i a and m o n o s p o r e - l i b e r a t i o n i n t h e C o n c h o c e l i s - p h a s e o f Por p h y r a t e n e r a K j e l l m . [ i n Japanese, E n g l i s h summary]. B u l l .  222  Tohoku Reg.  Fish.  Res.  Lab.  8:  45-61.  K u r o g i , fi. , and S . „ S a t o . 1962a. I n f l u e n c e s o f l i g h t on t h e g r o w t h and m a t u r a t i o n of C o n c h o c e l i s - t h a l l u s o f P o r p h y r a . I I . E f f e c t o f d i f f e r e n t p h o t o p e r i o d s on t h e growth and m a t u r a t i o n o f C o n c h o c e l i s - t h a l l u s o f P. t e n e r a K j e l l m a n [ i n J a p a n e s e , E n g l i s h summary]. B u l l . Tohoku Beg. f i s h . Res. Lab. 20: 127-137. K u r o g i , M., and S. S a t o . 1962b. I n f l u e n c e s o f l i g h t on t h e growth and m a t u r a t i o n o f C o n c h o c e l i s - t h a l l u s o f P o r p h y r a . i l l . E f f e c t of photoperiod i n the d i f f e r e n t species [ i n J a p a n e s e , E n g l i s h summary]. B u l l . Tohoku Reg. F i s h . B e s . Lab. 20: 138-156. K u r o g i , B . , and S. S a t o . 1967. E f f e c t o f p h o t o p e r i o d cn t h e g r o w t h and m a t u r a t i o n of C c n c h o c e l i s - t h a l l u s o f P o r p h y r a u m b i l i c a l i s <L.) K i i t z . and P o r p h y r a p s e u d o c r a s s a Yamada e t Mikami [ i n J a p a n e s e , e n g l i s h summary]. B u l l . T o h o k u Reg. F i s h . Bes. L a b . 27: 111-130. K u r o g i , 1 3 . , K. Akiyama and S. S a t o . 1962. I n f l u e n c e s o f l i g h t on t h e growth and m a t u r a t i o n o f C o n c h o c e l i s - t h a l l u s of P p r p h y r a . I . E f f e c t o f p h o t o p e r i o d on the f o r m a t i o n o f m o n o s p o r a n g i a and l i b e r a t i o n o f monospores [ i n J a p a n e s e , E n g l i s h summary]. B u l l . Tohoku Reg. F i s h . Res. Lab. 20: 121-126. K u r o g i , M., S, S a t o and T. Y o s h i d a . 1967. E f f e c t o f w a t e r t e m p e r a t u r e on t h e l i b e r a t i o n o f monospores from t h e C o n c h o c e l i s of P o r p h y r a u m b i l i c a l i s (L.) K i i t z . [ i n J a p a n e s e , E n g l i s h summary]. B u l l . Tohoku Beg. F i s h . Bes. Lab. 27: 131-139. K u s a k a b e , D. 1929. On t h e b i o l o g y and c u l t u r e o f t h e marine a l g a , Porphyra t e n e l l a [ t e n e r a ] K j e l l m . (Asakusanori). J . Imp. F i s h . I n s t . (Japan) 25: 21-22. K y l i n , B. .1922./deber d i e e n t w i c k l u n g s g e s c h i c h t e B a n g i a c e e n . A r k i v . B e t . 17: 1-12.  der  K y l i n , H. 1925. The m a r i n e r e d a l g a e i n t h e v i c i n i t y o f t h e b i o l o g i c a l s t a t i o n a t F r i d a y H a r b o r , Wash. Lunds O n i v e r s i t e t s A r s s k r i f t . N. F. Avd. 2, 2 1 ( 9 ) : 1-87. K y l i n , H. 1941. Irsskrift. Lee,  C a l i f o r n i s c h e Hhodophyceen. Lunds O n i v e r s i t e t s N. F. Avd. 2, 3 7 ( 2 ) : 1-51; p i s . 1-13.  B. E., and S. A. F u l t z . 1970. U l t r a s t r u c t u r e o f t h e c o n c h o c e l i s s t a g e o f t h e marine r e d a l g a P o r p h y r a l e u c o s t i c t a . J . P h y c o l . 6: 22-28.  L e w i n , J . 1966. S i l i c o n m e t a b o l i s m dioxide, a s p e c i f i c inhibitor 6: 1-12.  i n d i a t o m s . , V . Germanium of diatom growth. P h y c o l o g i a  223  Lin,  H.-P., M. fi. S o m m e r f e l d and J . R . S t r a f f o r d . 1975. L i g h t and e l e c t r o n m i c r o s c o p e o b s e r v a t i o n s on m o t i l e c e l l s o f PorphyrAdium purpureum ( R h o d o p h y t a ) . J . P h y c o l . 11: 452-457.  L i n d s t r o m , S. C, 1977. An A n n o t a t e d B i b l i o g r a p h y o f t h e B e n t h i c M a r i n e A l g a e o f A l a s k a . A l a s k a D e p t . o f F i s h and Game T e c h . D a t a R e p t . No. 3 1 ; i+172 pp. J u n e a u , A l a s k a . List,  R. J . 1966. S m i t h s o n i a n M e t e o r o l o g i c a l T a b l e s . S m i t h s o n i a n M i s c . C o l l e c t . 114: xi+527 pp., P u b l . 4014. S m i t h s o n i a n I n s t . , W a s h i n g t o n .  No.  L i i n i n g , K., K. S c h o i t z and J . W i l l e n b r i n k . 1972. T r a n s l o c a t i o n o f * C - l a b e l l e d a s s i m i l a t e s i n two Lamina r-i a s p e c i e s . pp. 420-425 i n : K. N i s i z a w a , e d . , P r o c . 7 t h I n t . Seaweed Symp., S a p p o r o , J a p a n 1971. U n i v . Tokyo P r e s s , T o k y o . 1  Magne, F. 1952. L a s t r u c t u r e du noyau e t l e c y c l e n u c l e a i r e c h e z l e P o r p h y r a l i n e a r i s G r e v i l l e . C. R. Acad. S c i . S e r . D, 234: 986-988. Magne, F. 1960. L e R h o d o c h a e t e p a r v u l a T h u r e t ( B a n g i o i d e e ) e t sa r e p r o d u c t i o n s e x u e e . C a h . B i o l . Mar. 1: 407-420. Markham, J . I . , a n d J . L . C e l e s t i n o . 1976. I n t e r t i d a l m a r i n e p l a n t s o f C l a t s o p C o u n t y , O r e g o n . S y e s i s 9: 253-266 [ I s s u e d May, 1 9 7 7 ] , Matsumoto, F, 1959. S t u d i e s on t h e e f f e c t o f e n v i r o n m e n t a l f a c t o r s on t h e growth o f * K o r i * ( P o r p h y r a t e n e r a K j e l l m . ) , w i t h s p e c i a l r e f e r e n c e t o t h e water c u r r e n t [ i n J a p a n e s e , E n g l i s h summary]. J . F a c . F i s h . Anim. Husb. H i r o s h i m a U n i v . 2: 249-333. M c B r i d e , D. .,L. 1972. S t u d i e s on some B r i t i s h C o l u m b i a n r e p r e s e n t a t i v e s o f t h e E r y t h r o p e l t i d a c e a e (Rhodophyceae, Bangiophycidae). Ph.D. t h e s i s . U n i v e r s i t y o f B r i t i s h C o l u m b i a , V a n c o u v e r , x i + 1 0 8 pp., p i s . 1-40. M c B r i d e , D. L . , and K. C c l e . 1969. U l t r a s t r u c t u r a l c h a r a c t e r i s t i c s o f t h e v e g e t a t i v e c e l l o f S m i t h o r a naiadum ( R h o d o p h y t a ) . P h y c o l o g i a 8: 177-186. M c E r i d e , D. L . , and K. C c l e . 1971. E l e c t r o n m i c r o s c o p i c o b s e r v a t i o n s on t h e d i f f e r e n t i a t i o n and r e l e a s e o f monospores i n t h e m a r i n e r e d a l g a S m i t h o r a naiadum. P h y c o l o g i a 10: 49-61. M c D o n a l d , K. L . 1972. L i f e - h i s t c r y a n d c y t o l o g i c a l s t u d i e s o f some Rhodophyceae. Ph.D. t h e s i s , U n i v e r s i t y o f C a l i f o r n i a , B e r k e l e y , vi+170 pp., f i g s . 1-83. M c L a c h l a n , J . , and J . S. C r a i g i e . 1966. A n t i a l g a l some s i m p l e p h e n o l s . J . P h y c o l . 2: 133-135.  a c t i v i t y of  224  Meer, J . P. van d e r . 1977. G e n e t i c s o f G r a c i l a r i a s p . (Rhodophyceae, G i g a r t i n a l e s ) I I . The l i f e h i s t o r y and genetic implications of cytokinetic f a i l u r e during t e t r a s p o r e f o r m a t i o n . P h y c o l o g i a 16; 367-371 . B e e r , J . P. van d e r , and E . R. Todd. 1977. G e n e t i c s o f G r a c i l a r i a s p . (Rhodophyceae, G i g a r t i n a l e s ) . I V . . - H i t o t i c r e c o m b i n a t i o n and i t s r e l a t i o n s h i p t o mixed phases i n t h e l i f e h i s t o r y . Can. J . B o t . 5 5 : 2810-2817. M i g i t a , S. 1959a. On t h e i n f l u e n c e o f e n v i r o n m e n t a l f a c t o r s on growth o f C o n c h o c e l i s and t h e c h a n g e i n pH o f c u l t u r e s e a water f i n J a p a n e s e , E n g l i s h summary ]. B u l l . Fac. , F i s h . N a g a s a k i U n i v . 8: 207-215. M i g i t a , S. 1959b. The i n v e s t i g a t i o n o f t h e C o n c h o c e l i s i n flriake S e a . I . The d i s t r i b u t i o n of s u b s t r a t u m a n d t h e p e r f o r a t i n g growth of c a r p o s p o r e s o f P o r p h v r a o f f Shimabara [ i n J a p a n e s e , E n g l i s h summary]. B u l l . F a c . F i s h , N a g a s a k i U n i v . 8: 216-222. M i g i t a , S. 1961. S t u d i e s on • P l a n t l e t o f C o n c h o c e l i s - p h a s e o f P o r p h y r a [ i n J a p a n e s e , E n g l i s h summary]. B u l l . F a c . F i s h . N a g a s a k i U n i v . 11: 128-136; p i s . 22-25. 1  M i g i t a , S. 1962. S t u d i e s on p l a n t l e t s o f C p n c f a c c e l j s - p h a s e o f P o r p h y r a . R e c o r d s c f O c e a n c g r . H o r k s J a p a n , S p e c i a l No. 6: 147-152. M i g i t a , S. 1964. F r e e z e - p r e s e r v a t i o n of P o r p h y r a t h a l l i i n v i a b l e s t a t e - I . V i a b i l i t y o f P o r p h y r a t e n e r a p r e se r v e d a t low t e m p e r a t u r e a f t e r f r e e z i n g i n t h e s e a water and f r e e z i n g under h a l f - d r i e d c o n d i t i o n [ i n J a p a n e s e , English summary]. B u l l . F a c . F i s h . N a g a s a k i Univ. 17: 44-54. M i g i t a , S. 1966. F r e e z e - p r e s e r v a t i o n of P o r p h y r a t h a l l i i n v i a b l e s t a t e - I I . E f f e c t o f c o o l i n g v e l o c i t y and water c o n t e n t o f t h a l l i on t h e f r o s t - r e s i s t a n c e [ i n J a p a n e s e , E n g l i s h summary]. E u l l . F a c . F i s h . N a g a s a k i U n i v . 21: 131-13 8. M i g i t a , S. 1967a. V i a b i l i t y and s p o r e - l i b e r a t i o n o f Conchocelis-phase, Porphyra t e n e r a , f r e e z e - p r e s e r v e d i n s e a water [ i n J a p a n e s e , E n g l i s h summary ]. B u l l . F a c . , F i s h . N a g a s a k i U n i v . 22: 33-43. M i g i t a , S. 1967b. C y t o l o g i c a l s t u d i e s on P o r p h y r a y e z o e n s i s Ueda, B u l l . F a c . F i s h . N a g a s a k i U n i v . 24; 55-64. M i g i t a , S. 1969. O l c l d i o p s i s d i s e a s e o f c u l t u r e P o r p h y r a f i n J a p a n e s e , E n g l i s h summary]. E u l l . F a c . F i s h . N a g a s a k i U n i v . 28: 131-145. M i g i t a , S. 1972. S t u d i e s on a t t a c h m e n t o f c c n c h c s p o r e and moncspcre o f P o r p h y r a y e z o e n s i s f i n J a p a n e s e , E n g l i s h summary]. B u l l . F a c . F i s h . N a g a s a k i U n i v . 33: 39-48.  225  M i g i t a , S. 1973. C h y t r i d d i s e a s e o f C o n c h o c e l i s i n P o r p h y r a c u l t i v a t i o n [ i n J a p a n e s e , E n g l i s h summary]. B u l l . F a c . F i s h . N a g a s a k i U n i v . 35: 41-48. M i g i t a , S. 1974. Some o b s e r v a t i o n s on f o r m a t i o n and l i b e r a t i o n of c o n c h o s p o r e s i n C o n c h o c e l i s o f P o r p h y r a [ i n J a p a n e s e , E n g l i s h summary]. B u l l . F a c , F i s h . N a g a s a k i O n i v . 38: 77-85. M i g i t a , S., and N, Abe. 1966, F o r m a t i o n o f s p o r e s i n C o n c h o c e l i s o f P p r p h y r a [ i n J a p a n e s e , E n g l i s h summary]. B u l l . F a c . F i s h . N a g a s a k i O n i v . 20: 1-13. M i g i t a , S., and C. B. Kim. 1970. S t u d i e s on h o r i z o n t a l growth o f C o n c h o c e l i s [ i n J a p a n e s e , E n g l i s h summary]. B u l l . F a c . F i s h . N a g a s a k i O n i v . 30: 1-8. M i g i t a , S., M. l a m a g u c h i and S. Tanaka. 1971. F r e e z e - p r e s e r v a t i o n o f germinated conchospores c f Porchyra y e z o e n s i s i n v i a b l e s t a t e [ i n J a p a n e s e , E n g l i s h summary]. B u l l . F a c . F i s h . N a g a s a k i O n i v . 31: 69-76. M i u r a , A. 1961. A new s p e c i e s o f P o r p h y r a and i t s C o n c h o c e l i s - p h a s e i n n a t u r e . J . Tokyo O n i v . F i s h . 305-311.  47:  M i u r a , A. 1968. P o r p h v r a k a t a d a i . a new s p e c i e s f r o m J a p a n e s e c o a s t . J . T o k y o O n i v . F i s h . 54: 55-59; p i s . 1-7. M i u r a , A. 1975. P o r p h y r a c u l t i v a t i o n i n Japan, pp. 273-304 i n : J . T o k i d a and H. H i r o y u k i , e d s . . Advance c f P h y c o l o g y i n J a p a n . Junk, The Hague. M i u r a , A., and S. I t o . 1959. An i n v e s t i g a t i o n o f C o n c h o c e l i s i n n a t u r e [ i n J a p a n e s e , E n g l i s h summary]. B u l l . J a p . S o c . P h y c o l . 7: 19-26. Mumford, T. F. J r . 1973a. O b s e r v a t i o n s on t h e taxonomy and e c o l o g y o f some s p e c i e s o f P o r p h v r a from W a s h i n g t o n and V a n c o u v e r I s l a n d , B r i t i s h C o l u m b i a . Ph.D. thesis. U n i v e r s i t y o f W a s h i n g t o n , S e a t t l e . x*164 pp., p i s . 1-38. Mumford, T. F. J r . 1973b. A new s p e c i e s o f P o r p h v r a f r o m west c o a s t c f N o r t h A m e r i c a . S y e s i s 6: 239-2 42.  the  Mumford, T i F. J r . 1975. O b s e r v a t i o n s on t h e d i s t r i b u t i o n and seasonal occurrence of Porphyra s c h i z o p h y l l a Hollenberg. P o r p h y r a t o r t a K r i s h n a m u r t h y , and P p r p h y r a b r u m a l i s s p . nov. ( B h o d o p h y t a , B a n g i a l e s ) . S y e s i s 8: 321-332 [ I s s u e d A p r i l , 1976], Mumford, T. F. J r . , and K. C o l e . 1977, Chromosome numbers f o r f i f t e e n s p e c i e s i n t h e genus P o r p h y r a ( B a n g i a l e s . Bhodophyta) from t h e west c o a s t o f N o r t h A m e r i c a . P h y c o l o g i a 16: 373-377.  226  N a g a i , M. 1941. M a r i n e A l g a e o f A g r i c . H o k k a i d o Ifflp. O n i v ,  the 46:  Kurile Islands. II. J . 139-310; p i s . 4-6.  N a t i o n a l R e s e a r c h C o u n c i l o f Canada, A s t r o p h y s i c s Branch, D a i l y s u n r i s e and s u n s e t t a b l e s (1974) f o r V i c t o r i a ,  Fac. 1974. B.C.  N i c h c l s , H. W., and E, L i s s a n t . 1967. D e v e l o p m e n t a l s t u d i e s o f E r y t h r o c l a d i a R o s e n v i n g e i n c u l t u r e . J . P h y c o l . 3: 6-18. O g a t a , E. 1955. P e r f o r a t i n g growth o f C o n c h o c e l i s m a t r i c e s , Bot. Mag. l o k y o 68: 371-372. O g a t a , E. 1959. O b s e r v a t i o n s on t h e v e r t i c a l C o n c h o c e l i s . B o t . Mag. T o k y o 72: 177*  in  growth  calcareous  of  O g a t a , £. 1961. S t u d i e s on t h e qrowth o f C o n c h o c e l l s f i n J a p a n e s e , E n g l i s h summary ]. J . S h i m o n o s e k i C o l l e g e o f F i s h e r i e s 10: 69-146; p i s . 1-12. O g a t a , E . 1963. M a n o m e t r i c s t u d i e s on t h e r e s p i r a t i o n o f a marine a l g a , P o r p h y r a t e n e r a - I , I n f l u e n c e of s a l t c o n c e n t r a t i o n , t e m p e r a t u r e , d r y i n g and o t h e r f a c t o r s [ i n J a p a n e s e , E n g l i s h summary]. B u l l . J a p . Soc. S c i . F i s h . 29: 139-145., O g a t a , E. 1971. Growth o f c o n c h o c e l i s i n a r t i f i c i a l medium i n r e l a t i o n t o c a r b o n d i o x i d e and c a l c i u m m e t a b o l i s m . J , S h i m c n e s e k i O n i v . F i s h . 19: 59-65. O g a t a , E . 1975. P h y s i o l o g y o f P o r p h y r a . pp. 151-160 i n : J . T o k i d a and H. H i r o y u k i , e d s . . A d v a n c e o f P h y c o l o g y i n J a p a n . J u n k , The Hague. O g a t a , E., and w. Schramm. 1971. Some o b s e r v a t i o n s on t h e i n f l u e n c e o f s a l i n i t y on g r o w t h and p h o t o s y n t h e s i s i n P o r p h y r a u m b i l i c a l i s . Mar. B i o l . 10: 7 0-76. Ohmi, H. 1963. On e p i p h y t i c s p e c i e s o f P o r p h y r a and t h e i r h o s t s [ i n J a p a n e s e , E n g l i s h summary and t a b l e ]. B u l l . J a p . Soc. P h y c o l . 11: 38-44. Okamura, K., K. Onda and M. H i g a s h i . 1920. P r e l i m i n a r y n o t e s the development o f the c a r p o s p o r e s of Pprphyra tenara C t e n e r a ] K j e l l m . B o t . Mag. Tokyo 34: 131-135; p l . 3.  on  P a p e n f u s s , G. F. 1955. C l a s s i f i c a t i o n o f t h e A l g a e , pp. 115-224 i n : E. L. K e s s e l , e d . , A C e n t u r y o f P r o g r e s s i n t h e N a t u r a l S c i e n c e s 1853-1953. C a l i f . Acad. S c i . , San Francisco. P a r k e r , B. C. 1966. T r a n s l o c a t i o n i n M a c r o c y s t i s , I I I . C o m p o s i t i o n o f s i e v e t u b e e x u d a t e and i d e n t i f i c a t i o n o f t h e major C * - l a b e l l e d p r o d u c t s . J . P h y c o l , 2: 38-41, 1  P a r s o n s , T, B, governing  1965, A g e n e r a l d e s c r i p t i o n c f some f a c t o r s primary production i n the S t r a i t of G e o r g i a ,  227  H e c a t e S t r a i t and Queen C h a r l o t t e Sound, and t h e N. I . P a c i f i c Ocean. F i s h , R e s . B o a r d Canada, M a n u s c r i p t R e p t . . S e r i e s ( O c e a n o g r a p h i c and l i m n o l o g i c a l ) No. 193: 1-34; f i g s . 1-13, t a b l e s 1-7. P e y r i e r e , M. 1971. E t u d e i n f r a s f r u c t u r a l e des G r i f f i t h s j , a f l o s c u l p s a ( R h o d c p h y c e e ) . C. S e r . D, 273: 2071-2074.  spermatocystes R. A c a d . S c i .  du  P e y r i e r e , M. 1974. Etude i n f r a s f r u c t u r a l e des £.permaties de d i f f e r e n t e s R h c d o p h y c e e s F l o r i d e ' e s . C. , R. Acad. S c i . S e r . D, 278: 1019-1022. P r i n g l e , J . D. , and A. P. A u s t i n . 1970. The m i t o t i c i n d e x i n selected red algae i n s i t u . I I . A s u p r a l i t t o r a l species, P o r p h y r a l a n c e o i a t a ( S e t c h e l l S Hus) G. M. S m i t h , J . Exp, M a r . ~ B i o l 7 E c o l . 5: 113-137. P r o v a s o l i , L. 1971. M e d i a and p r o s p e c t s f o r t h e c u l t i v a t i o n o f marine a l g a e , pp. 599-604 i n : J . Rosowski and B. P a r k e r , eds., S e l e c t e d Papers i n P h y c o l o g y . , U n i v e r s i t y of Nebraska, L i n c o l n . Prud'homme van R e i n e , 8. F., and C. van den Hoek. 1966. I s o l a t i o n o f l i v i n g a l g a e g r o w i n g i n t h e s h e l l s of m c l l u s c s and b a r n a c l e s w i t h EDTA ( E t h y l e n e d i a m i n e t e t r a a c e t i c A c i d ) . Blumea 14: 331-332. Ramus, J . 1969. The d e v e l o p m e n t a l s e g u e n c e o f t h e m a r i n e r e d a l g a P s e u d o g l o i o p h l o e a i n c u l t u r e . Univ. C a l i f . B e r k e l e y P u b l . B o t . 52: 1-427 B e e s , T. K. 1940. A p r e l i m i n a r y a c c o u n t P p r p h y r a u m b i l i c a l i s (L.) Ag. Ann. 669-671.  o f the l i f e - h i s t o r y Bot. (London) N. S.  B e n t s c h l e r , H.-G. 1967, P h o t o p e r i c d i s c h e i n d u k t i o n d e r monosporenbildung b e i Porphyra t e n e r a K j e l l m . P l a n t a 65-74 R e y n o l d s , E. S. 1963. The u s e o f l e a d c i t r a t e a t h i g h e l e c t r o n opague s t a i n i n e l e c t r o n m i c r o s c o p y . , J . B i o l . 17: 208-212. R i c h a r d s o n , N. 1970. S t u d i e s on t h e p h o t o b i o l o g y o f f u s c o p u r p u r e a . J . P h y c o l . 6: 215-219. B i c h a r d s o n , N. 1972. Spore c l a s s i f i c a t i o n i n t h e P o r p h y r a . B r . P h y c o l . J . 7: 49-51.  of 4:  76:  pfl as Cell.  an  Ban-gia  genera  Bangia  B i c h a r d s o n , N., and P. S. D i x o n . 1968. L i f e h i s t o r y o f B a n g i a f u s c o p u r p u r e a ( D i l l w . ) L y n g t . i n c u l t u r e . N a t u r e (London) 218: 496-497. t  B o s e n v i n g e , I . K. 1909. The m a r i n e a l g a e o f Denmark, e t c . . P a r t I . I n t r o d u c t i o n . Rhodophyceae I . ( B a n g i a l e s and  228  N e m a l i o n a l e s ) . K• Dan, V i d e n s . S e l s k . S k r i f t e r 7, N a t u r v i d e n s k . Math, A f d . 7 ( 1 ) ; 1-151; p i s . 1-2, 2 maps. R o s e n v i n g e , 1. K. 1927. On m o b i l i t y i n t h e r e p r o d u c t i v e o f t h e R h o d o p h y c e a e . B o t . T i d s s k r . 40: 72-80.  cells  R o s e n v i n g e , L. K. 1931. The m a r i n e a l g a e o f Denmark. P a r t IV. Bhodophyceae IV ( G i g a r t i n a l e s , R h o d y m e n i a l e s , N e m a s t c m a l e s ) . K. Dan. V i d e n s k . S e l s k . S k r i f t e r 7, N a t u r v i d e n s k . Math. A f d . ..7 ( 4 ) : 492-627; p l . 8. S a i t o , Y. 1955. On t h e r e l a t i o n between t h e a p p e a r a n c e o f s p o r e s o f the l a v e r ( P o r p h y r a t e n e r a ) and t h e environmental c o n d i t i o n s - I [ i n Japanese, E n g l i s h summary]. E u l l . J a p . S o c . S c i . F i s h . 21: 226-228. S a i t o , Y. 1956. On the l i b e r a t i o n o f s p o r e s f r o m t h e C o n c h o c e l i s o f P o r p h y r a t e n e r a i n t h e sea ( P r e l i m i n a r y r e p o r t ) [ i n J a p a n e s e , E n g l i s h summary]. B u l l . J a p , S e c . S c i . F i s h . 21: 1215-1218. S a k u r a i , Y., K..Akiyama and S. S a t o . 1974. On t h e f o r m a t i o n and the d i s c h a r g e c f z o o s p o r e s o f Pythium porphyrae i n e x p e r i m e n t a l c o n d i t i o n s [ i n J a p a n e s e , E n g l i s h summary]. B u l l . Tohoku Beg. F i s h . R e s . L a b . 33: 119-127. S a s a k i , M., and Y. S a k u r a i . 1972. C o m p a r a t i v e o b s e r v a t i o n s on t h e growth among t h e f i v e s t r a i n s i n P y t h i u m porpfrvrae under t h e same c u l t u r a l c o n d i t i o n [ i n J a p a n e s e , E n g l i s h summary]. B u l l . Tohoku Reg. F i s h . Res. Lab. 32: 83-87. S a s a k i , M., and S. S a t o . 1969. C o m p o s i t i o n o f medium and c u l t u r a l temperature o f Pythium sp., a p a t h o g e n i c fungus, o f t h e " A k a g u s a r e " d i s e a s e o f c u l t i v a t e d P o r p h y r a Tin J a p a n e s e , E n g l i s h summary]. B u l l . Tohoku Reg. F i s h . Res. Lab. 29: 125-132. S c a g e l , B. F. 1957. An A n n o t a t e d L i s t o f t h e M a r i n e A l g a e o f B r i t i s h C o l u m b i a and N o r t h e r n S a s h i n g t c n . Nat. Museum o f Canada B u l l . Nc. 150: vi+289 pp. Ottawa. S c a g e l , R. F. 1973. M a r i n e b e n t h i c p l a n t s i n t h e v i c i n i t y c f B a m f i e l d , B a r k l e y Sound, B r i t i s h C o l u m b i a . S y e s i s 6: 127-145. S c h l i c h t i n g , H. E . J r . 1975a. B i o m o n i t o r t e c h n i g u e s f o r b i o a s s a y i n g w a t e r g u a l i t y . B i o c o n t r o l T e c h n i g u e s 1: S c h l i c h t i n g , H. E . J r . 1975b. The b i o m o n i t o r . C a r c l i n a 3 8 ( 9 ) : 1 p . C a r o l i n a B i o l . S u p p l y Co.  4-5.  Tips  S c h l i c h t i n g , H. E. J r . 1976. B i o m o n i t o r l a b o r a t o r y e x e r c i s e s . C a r c l i n a T i p s 3 9 ( 4 ) ; 13-14. C a r o l i n a B i o l . S u p p l y Co. S c o t t , J . L., and P. S. D i x o n . 1 9 7 3 a . , O l t r a s t r u c t u r e o f t e t r a s p c r o g e n e s i s i n the marine red a l g a P t i l o t a  229  hypnpides. J . Phyccl.  9:  2 9-46.  S c o t t , J . L . , and P. S. D i x o n . 1973b. U l t r a s t r u c t u r e o f spermatium l i b e r a t i o n i n t h e marine r e d a l g a P t i l o t a d e n s a . J . P h y c o l . S: 85-91, Setchell, and N . L . America. Univ. C a l i f . p i s . 17-27.  G a r d n e r . 1903. A l g a e o f n o r t h w e s t e r n B e r k e l e y , P u b l . B o t . 1: 165-418;  Sharma, A. K., and A. Sharma. 1965. Chromosome T e c h n i g u e s ; t h e o r y and P r a c t i c e . B u t t e r w c r t h s and Co. L t d . , London, x+474 pp. S h e a t h , R. G., J . A. H e l l e b u s t and T. Sawa. 1977. C h a n g e s i n p l a s t i d s t r u c t u r e , p i g m e n t a t i o n and p h o t o s y n t h e s i s o f t h e c o n c h o c e l i s s t a g e o f fforphyra l e u c o s t i c j t a (Rhodophyta, B a n g i o p h y c e a e ) i n r e s p o n s e t o low l i g h t and d a r k n e s s . P h y c o l o g i a 16: 265-276. S h i n m u r a , I..1974. P o r p h y r a t a n e q a s h i m e n s i s . a new s p e c i e s o f Rhodophyceae f r o m T a n e g a s h i m a I s l a n d i n s o u t h e r n J a p a n . B u l l . J a p . Soc. S c i . F i s h . 40: 735-749. S i e b u r t h , J . McN., and A. J e n s e n . 1969. S t u d i e s on a l g a l s u b s t a n c e s i n t h e s e a . I I . The f o r m a t i o n o f G e l t s t o f f (humic m a t e r i a l ) by e x u d a t e s o f P h a e o p h y t a . J . E x p . Mar. B i o l . E c o l . 3: 275-289. S i m o n - B i c h a r d - B r e a u d , J . 1971. On a p p a r e i l c i n e t i q u e dans l e s g a m e t o c y s t e s maTes d'une Rhodophycee; B o n n e m a i s c n i a h a m i f e r a H a r i o t . C. R. A c a d . S c i . S e r . D, 273: 1272-1275; p i s . T-4. S i m o n - B i c h a r d - B r e a u d , J . 1972a. O r i g i n e e t d e v e n i r d e s v a c u o l e s \ p o l y s a c c h a r i d e s des g a m e t o c y s t e s males de B o n n e m a i s c n i a h a m i f e r a H a r i o t ( R h o d o p h y c e e ) . C. R. A c a d . S c i . S e r . D, 274:~1485-1488; p i s . 1-4. S i m c n - B i c h a r d - B r e a u d , J . .1972b* F o r m a t i o n de l a c r y p t e f l a g e l l a i r e e t e v o l u t i o n de son c o n t e n u a u c o u r s de l a gametogenese maMe c h e z B o n n e m a i s o n i a h a m i f e r a H a r i o t ( R h o d o p h y c e e ) . , C . R. A c a d . S c i . S e r . D, 274: 1796-1799; pis. 1-4. S m i t h , G. M. 1944. M a r i n e A l g a e o f t h e M o n t e r e y P e n i n s u l a C a l i f o r n i a . S t a n f o r d U n i v . P r e s s , S t a n f o r d , ix+622 pp; pis. 1-98. S m i t h , G. M., and G. J . H o l l e n b e r g . 1943. On some Rhodophyceae from t h e M o n t e r e y P e n i n s u l a , C a l i f o r n i a . Am. J . B o t . 30; 211-222. Sommerfeld, M.R., and H. H. N i c h o l s . 1970. D e v e l o p m e n t a l and c y t o l o g i c a l s t u d i e s of Bangia fuscopurpurea i n c u l t u r e . Am. J . B o t . 57: 640-64 8.  230  S c m m e r f e l d , fl. B., and H.W. N i c h o l s . 1973. The l i f e c y c l e o f B a n g i a f u s c o p u r p u r e a i n c u l t u r e . I . E f f e c t s of t e m p e r a t u r e and p h o t o p e r i o d on t h e m o r p h o l o g y and r e p r o d u c t i o n c f t h e B a n g i a phase. J . P h y c o l . 9: 205-210. , S p a r l i n g . S. B. 1971. B e c e n t r e c o r d s o f marine a l g a e i n San L u i s O b i s p o C o u n t y , C a l i f o r n i a . P h y c o l o g i a 10: 235-210. S p a r r o w , F. K. 1969. Z o o s p o r i c marine f u n g i from t h e P a c i f i c n o r t h west (U.S.A.). A r c h . M i k r o b i o l . 66: 129-146. S t e g e n g a , H,, and » . J . B o r s j e . 1976. The m o r p h o l o g y and h i s t o r y o f Acrochaetium dasyae C o l l i n s (Bhodophyta, N e m a l i a l e s ) . A c t a B o t . N e e r l . 24: 15-29.  life  S t e g e n g a , 8., and M. Vroman. 1976. The morphology and l i f e h i s t o r y o f A c r o c h a e t i u m densum (Drew) P a p e n f u s s ( B h o d o p h y t a , N e m a l i a l e s ) . A c t a B o t . N e e r l . 25: 257-280. S t c s c h , fi. A. von. 1965. The s p o r o p h y t e o f L i a g p r a f a r i n o s a Lamour. B r . P h y c o l . B u l l . 2: 486-496. S u t o , S. 1963. I n t e r g e n e r i c and i n t e r s p e c i f i c c r o s s i n g s o f t h e l a v e r s ( P o r p h y r a ) . B u l l . J a p . Soc. S c i . F i s h . 29: 739-748. S u t o , S., T. Maruyama and 0. U m e b a y a s h i . 1954. On t h e s h e d d i n g o f s p o r e s f r o m " c o n c h o c e l i s - p h a s e " o f Pgr p h y r a t e n e r a c u l t u r e d i n s h e l l s of b i v a l v e s [ i n Japanese, E n g l i s h summary]. ...Bull. J a p . S o c . S c i . F i s h . 20: 490-493. S v e d e l i u s , N. 1917. Die monosporen b e i H e Im i n t h o r a d-i var l e a t a n e b s t n o t i z iiber d i e z w e i k e r n i g k e i t i h r e s karpogons. Ber. D e u t s c h Bot. Ges. 35: 212-224. T a k a h a s h i , fl., T. I c h i t a n i and H. S a s a k i . 1977. Pythium p o r p h v r a e T a k a h a s h i jet S a s a k i , s p . nov. c a u s i n g r e d r o t c f m a r i n e r e d a l g a e P o r p h y r a s p p . T r a n s , flycol. S o c . J a p a n 18: 279-285. T a k e u c h i , T., T. H a t s u b a r a , fl. S h i t a n a k a and S. S u t o . 1954. On the s h e d d i n g o f s p c r e s from c u l t u r e d " c o n c h o c e l i s - p h a s e " °f P o r p h y r a t e n e r a s e t i n t h e s e a [ i n J a p a n e s e , E n g l i s h summary]. B u l l . J a p . S o c . S c i . F i s h . 20: 487-489. T a n a k a , K. 1959. A s t u d y on t h e c u l t u r e o f t h e c o n c h o c e l i s o f P g r p h y r a t e n e r a on t h e bottom o f t h e s e a [ i n J a p a n e s e , E n g l i s h summary]. B u l l . J a p , S o c . S c i . F i s h . 24: 701-704. T a n a k a , T. 1952. The s y s t e m a t i c s t u d y o f t h e J a p a n e s e P r o t o f l o r i d e a e . Mem. F a c . F i s h . , Kagoshima U n i v . 2 ( 2 ) : 1-92; p i s . 1-23. T a y l o r , «. B. 1945. P a c i f i c m a r i n e a l g a e o f t h e A l l a n Hancock e x p e d i t i o n s t o t h e G a l a p a g o s I s l a n d s . A l l a n Hancock P a c i f i c E x p e d . 12: i v + 5 2 8 .  231  T h u r e t , G., and E. B o r n e t . 1878. E t u d e s P h y c o l c q i q u e s . P a r i s , Masson. i i i * 1 0 5 pp; p i s . 1-51. T o k i d a , J . 1960. M a r i n e a l q a e e p i p h y t i c on l a m i n a r i a l e s B u l l . F a c . F i s h . H o k k a i d o O n i v . 11: 73-105.  plants.  T s e n q , C. K., and T. J . C h a n q . 1955. S t u d i e s on P o r p h y r a . I I I . Sexual reproduction i n Porphvra T i n Chinese, E n q l i s h summary]. A c t a B o t . S i n i c a 4: 153-166; p i s . 1-3. T s e n q , C. K., and T. J . Chanq. 1956. C o n d i t i o n s o f P o r p h v r a c o n c h o s p o r e s f o r m a t i o n and d i s c h a r q e and t h e d i s c h a r q e r h y t h m . A c t a B o t . S i n i c a 5: 33-48. T s e n q , C. K., T. J . C h a n q , and I. ¥, 2hao. 1963. C o m p a r a t i v e , s t u d i e s on t h e i n f l u e n c e o f t h e t e m p e r a t u r e f a c t o r o r t h e f o r m a t i c and d i s c h a r g e o f c o n c h o s p o r e s o f d i f f e r e n t s p e c i e s o f P o r p h y r a f i n C h i n e s e , E n q l i s h summary]. A c t a B o t . S i n i c a 11: 261-271. T s u k i d a t e , J . - I . 1970. M i c r o b i o l o q i c a l s t u d i e s o f P o r p h y r a p l a n t s - I . S t u d i e s o f b a c t e r i a i s o l a t i o n methods. B u l l . N a n s e i Req. F i s h . Bes. L a b . 3; 19-22. T s u k i d a t e , J . - I . 1971. M i c r o b i o l o q i c a l s t u d i e s o f P o r p h v r a p l a n t s - I I I . A b n o r m a l i t y on t h e qrowth o f P o r p h v r a p l a n t s by t h e d i s t u r b a n c e o f t h e b a c t e r i a l f l o r a a c c o m p a n y i n q t h e p l a n t . B u l l . N a n s e i Req. F i s h . R e s . Lab. 4: 1-12. T s u k i d a t e , J . - I , 1974. M i c r o b i o l o q i c a l s t u d i e s o f P o r p h y r a p l a n t s - I v . On t h e r e l a t i o n between t h e qrowth o f t h e P o r p h y r a p l a n t s c u l t i v a t e d i n t h e s e a and t h e b a c t e r i a l f l o r a accompanyinq t h e p l a n t s . B u l l . N a n s e i Beq. F i s h . Res. Lab. 7: 1-8. T s u r u q a , H., and T. N i t t a , 1960. B i o c h e m i c a l i n v e s t i q a t i c n s o f t h e marine a l q a e . I I I . D i n a m i c [ s i c ] c o r r e l a t i o n between h o s t and p a r a s i t e i n P y t h i u m d i s e a s e o f P o r p h y r a t e n e r a [ i n J a p a n e s e , E n q l i s h summary]. H i r o s h i m a N a k a i Req. F i s h . B e s . L a b . , B u l l . .13: 19-22. Ueda, S. 1929. On t h e l i f e - h i s t o r y J . Imp. F i s h . I n s t . T o k y o 24:  of Porphyra 139-142.  tenera Kjellm.  V o z z h i n s k a y a , V. B. 1964. M a k r o f i t y m o r s k i k h p o b e r e z h i y S a k h a l i n a {Macrophytes o f t h e S a k h a l i n s e a c o a s t ) . T r u d y I n s t . O k e a n o l . ,69: 330-441. [ i n R u s s i a n , E n q l i s h t r a n s l a t i o n p u b l i s h e d by U.S. N a v a l O c e a n o q r . O f f i c e , T r a n s l a t i o n No. 4 6 2 ] . w e t h e r b e e , R., and M. J . Wynne. 1973. The f i n e s t r u c t u r e o f t h e nuclear associations of developinq carpospcranqia i n P c l y s i p h o n i a n o v a e - a n g l i a e ( B h o d o p h y t a ) . J . P h y c o l . 9: 402-407. Hiddowson, T.  B.  1974.  The  marine  alqae o f B r i t i s h Columbia  and  232  n o r t h e r n W a s h i n g t o n : R e v i s e d l i s t and k e y s . P a r t . I I . B h o d o p h y c e a e ( r e d a l g a e ) . S y e s i s 7: 143-166 [ I s s u e d J a n . 1975]. Wittmann, W. 1965. A c e t o - i r o n - h a e m a t o x y l i n - c h l o r a l h y d r a t e f o r chromosome s t a i n i n g . S t a i n T e c h . 40: 161-164. Woessner, J . W. 1974. The measure and h a r v e s t o f t h e C a l i f o r n i a m a r i n e c r o p p l a n t P o r p h v r a . p. A61 i n : A b s t r a c t s o f P a p e r s , 8 t h I n t . Seaweed Symp., B a n g o r , W a l e s . Woessner, J . W., P. S o r e n s o n and D. Coon. 1977. The e c o n o m i c p o t e n t i a l o f h a r v e s t i n g and m a r k e t i n g an A m e r i c a n P o r p h y r a . J . P h y c o l . S u p p l . t o 13: 74 ( A b s t r . ) . Wynne, M. J . 1972. The genus P o r p h y r a a t A m c h i t k a I s l a n d , A l e u t i a n s , pp. 100-104 i n : K. N i s i z a w a , e d . , P r o c . 7 t h I n t . Seaweed Symp., S a p p o r o , J a p a n 1971. Tokyo O n i v . P r e s s , Tokyo. Y a i u , H. 1967. N u c l e a r d i v i s i o n i n B a n g i a f u s c o p u r p u r e a . ( B i l l w y n ) L y g b y e . E u l l . F a c . F i s h . , H o k k a i d o U n i v . 17: 163-164; p i s . 1-5. Yabu, H. 1969a. O b s e r v a t i o n s on chromosomes i n some s p e c i e s o f £2IEkYX§.. B u l l . F a c . F i s h . , H o k k a i d o U n i v . 19: 239-243. Yabu, H. 1969b. M i t o s i s i n P o r p h y r a t e n e r a K j e l l m . B u l l . F a c . F i s h . , H o k k a i d o O n i v . 20: 1-3; p l s . ~ 1 - 2 . Yabu, H. 1970. C y t o l o g y i n two s p e c i e s o f P o r p h y r a f r o m t h e s t i p e s o f N e r e o c y s t i s l u e t k e a n a (Mert. ) P o s t , e t Rupr. B u l l . F a c . F i s h . , H o k k a i d o Oniv. 20: 243-251. Yafcu, H. 1971. O b s e r v a t i o n s on chromosomes i n some s p e c i e s o f P o r p h y r a I I . B u l l . F a c . F i s h . , H o k k a i d o O n i v . 2 1 : 253-258; p i s . 1-9. Yabu, H. 1972. O b s e r v a t i o n s on chromosomes i n some s p e c i e s o f P o r p h v r a I I I . B u l l . F a c . F i s h . , H o k k a i d o O n i v . 22: 261-266. Yabu, H. 1975. C y t o l o g i c a l s t u d i e s i n t h e R h o d o p h y t a and C h l o r o p h y t a . pp. 125-135 i n : J . T o k i d a and H. H i r o s e , e d s , , Advance o f P h y c o l o g y i n J a p a n . J u n k , The Hague. Yabu, H., and J , T o k i d a . 1963. M i t o s i s i n P o r p h y r a . F i s h . , H o k k a i d o O n i v . 14: 131-136; p i s . 1-6.  B u l l . Fac,  Y a m a s a k i , H. 1954a. S t u d i e s on t h e e c o l o g y o f t h e conchocelis-phase of Porphyra tenera Kjellm. - I f i n J a p a n e s e , E n g l i s h summary]. B u l l . J a p . S o c . S c i . F i s h . 20; 4 42-446; p i s . 1. Y a m a s a k i , H. 1954b. S t u d i e s on t h e e c o l o g y o f t h e conchocelis-phase cf Porphyra tenera K j e l l m . - I I . ,  233  S h e d d i n g and f i x i n g o f s p o r e s [ i n J a p a n e s e , E n g l i s h summary]. B u l l . J a p . Soc. S c i . F i s h . 20: 447-450. Yendo, K. 1919. T h e g e r m i n a t i o n and d e v e l o p m e n t o f some a l g a e . I . B o t . Mag. Tokyo 3 3 : 73-93; p i . 1.  marine  Young, D. N. 1977. A n o t e on t h e a b s e n c e o f f l a g e l l a r s t r u c t u r e s i n s p e r m a t i a of Bonnemaisonia. P h y c o l o g i a 219-222.  16:  Z i n o v a , E. S. 1940. M o r s k i e v o d o r o s l i K o m a n d o r s k i k h O s t r o v o v [ i n R u s s i a n , E n g l i s h summary, 'The a l g a e c f t h e Commander I s l a n d s ' , pp. 2 3 6 - 2 3 7 ] . T r u d y T i k h o o k e a n s k o g o K o m i t e t a , L e n i n g r a d 5: 166-243. Z i n o v a , E. S. 1 9 5 4 . / V o d o r o s l i O k h o t s k o g o Morya [ i n R u s s i a n , t r a n s l a t e d t i t l e 'Algae o f t h e Okhotsk Sea']..Trudy B e t . I n s t . Akad. Hauk SSSB, s e r . 2, 9: 259-310.  234  APPENDIX I  Porphyra C o l l e c t i n g Jrifish  Stations  And, D a t e s Of C o l l e c t i o n s  Columbia:  Aguilar Point, 1977.  4 8 ° 50.4*N,  1 2 5 ° 08.3'W, 31 MA? 1974, 2 APR  B o l k u s I s l a n d s , S k i n c u t t l e I n l e t , Queen C h a r l o t t e 52° 19.4»N, 131° 14.8*W, 27 MAR 1976. Botanical  B e a c h , 4 8 ° 32*N,  Brooks P e n i n s u l a , Cable  124° 27*W, 11 J U L 1975.  5 0 ° 06.7* N, 1 2 5 ° 54.5*13, 25 JUN 1S75.  Beach, 4 8 ° 49.6'N,  Cape B e a l e , 4 8 ° 48 *N, Execution  125° 09.1»fl, 8 AUG 1974.  1 2 5 ° 13*W, 28 J U L 1976.  Rock, 4 8 ° 48.8*N,  1 2 5 ° 10.6*8, 10 MAY 1974.  G o r d o n I s l a n d s , Queen C h a r l o t t e 131° G8.7*W, 1 APR 1976. Grassy  Islands,  5 2 ° 06.1*N,  I s l a n d , K y u g u o t Sound, 49° 55. 4* N, 1 2 7 ° 15.0*8, 14 MAY 1975..  Hope I s l a n d , Hot  Islands,  Springs  5 0 ° 54»N, 127° 57' H, C o v e , 4 9 ° 21.5*N,  15 APR 1 975.  1 2 6 ° 15.8'«,  13 SAY 1975.  L a n g a r a I s l a n d , Queen C h a r l o t t e I s l a n d s , 5 4 ° 13.54*N, 132° 57.7 'W, 29 MAR 1S76.. L e a c h I s l e t , B a r k l e y Sound, 4 8 ° 49. 8* N, 1 2 5 ° 14.4*8, 2 APR 1977, 17 MAY 1977. R e n n e l Sound, Queen C h a r l o t t e I s l a n d s , 132° 30.7*8, 30 MAR 1S76. Ross I s l e t s , 1974.  Barkley  Second  Beach, B a r k l e y 1974.  South  Pender I s l a n d ,  Sound, 48° 52.2*N,  5 3 ° 22.65*N, 1 2 5 ° 0 9 . 5 » W , 5 JUN  Sound, 48° 48,9»N, 1 2 5 ° 1 0 « H , 9 MAY 48° 44,0*N,  123° 11.2«w, 4 SEP 1977.  Study s i t e , B a r k l e y Sound, 4 8 ° 50.1*N, 1974 t h r o u g h MAY 1977.  1 2 5 ° 11.1»W, JUN  T r e e I s l e t s , Queen C h a r l o t t e Sound, 5 0 ° 58.9*N, 127° 42.6'W, 24 JUN 1975.  235  V o l c a n i c C o v e , Kyuguot 15 MAY 1975. Hhiffen  Spit,  C h a n n e l , 4 9 ° 58.7*N, 127°  S o o k e , 48° 2 1 . 2 » N ,  123° 4 4 . 2 » W ,  13.9*»,  15 OCT  1S73.  Crejgcn: Harris  Beach,  42° 04»N,  124°  18*0, 23 SEP  1975.  California; Ano  Nuevo I s l a n d , 3 7 ° 0 6 » N ,  Mission  Point,  122° 20*W, 2 NOV  C a r m e l , 36° 32.5* N , 121° 56»H, 13 JUL  Pebble Beach, Monterey NOV 1975.  Peninsula,  1976.  36° 34*N, 121° 57*8,  Fescadero P o i n t , Monterey P e n i n s u l a , 12 J U L 1976., Feint  1975.  4  36° 33.8*N, 121° 5 7 » H ,  J o e , M o n t e r e y P e n i n s u l a , 36° 36.6*N, 121° 3 NOV 1975, 10 JUL 1976.  57.4*8,  236  APPENDIX  Representative specimens of porphyra a n d P. thuretii Porphyra  II  gardneri*  P.  nereocystis*  gardneri  Mcncsporic Alaska: Murder P o i n t , A t t u I s l a n d , 5 2 ° 4 8 ' N , 1 7 3 ° 11 « E , UBC 7 9 4 7 , 8 JUN 1 9 6 0 ; B i r d C a p e , A m c h i t k a I s l a n d , 5 1 ° 3 9 . 5 ' N , 1 7 8 ° 3 8 . 5 ' E , DBC 5 4 2 0 2 , 27 J U N 1 9 6 9 ; « T o p C a m p ' , A a c h i t k a I s l a n d , 5 1 ° 3 4 . 7 » N , 178<> 5 0 . 5 « E , UBC 5 4 2 0 3 , 17 AUG 1 9 7 0 ; C a p e A g a g d a k , A d a k I s l a n d , 5 2 ° 0 0 » N , 1 7 6 ° 3 5 « H , UBC 7 7 8 3 , 12 J U N 1 9 6 0 ; H a r b o r P o i n t , L i t u j a B a y , S i t k a , 5 7 ° C 8 » N , 1 3 5 ° 1 5 » 8 , UBC 2 3 4 0 1 , 29 J U N 1 9 6 5 ; C a p e H u z o n , D a l l I s l a n d , 5 4 ° 4 0 » N , 1 3 2 ° 4 1 » H , U E C 2 1 4 6 9 , 13 J U N 1 9 6 5 . British  Columbia:  L a n g a r a I s l a n d , Queen C h a r l o t t e I s l a n d s , 5 4 ° 1 3 . 5 4 * N , 1 3 2 ° 57.7*W, U B C 5 4 8 8 3 , 29 MAR 1 9 7 6 ; P l o v e r I s l a n d , 5 0 ° 5 5 ' N , 1 2 7 ° 5 8 « B , UBC 1 5 0 1 , 2 6 J U N 1 9 5 3 ; G a r d e n I s l a n d , K y u g u o t , 5 0 ° 0 1 « N , 1 2 7 ° 2 1 » W , UBC 4 3 4 2 , 2 7 MAY 1 9 5 9 ; D i a n a I s l a n d , B a r k l e y S o u n d , 4 8 ° 5 0 . 1 » N , 1 2 5 ° 1 1 . 1 * 8 , UBC 5 4 1 3 3 , 2 3 F E B 1 9 7 5 ; UBC 5 4 1 2 7 a n d UBC 5 4 1 3 4 , 28 MAR 1 9 7 5 ; UBC 5 4 8 1 9 , 16 APR 1 S 7 6 ; E x e c u t i o n R o c k , B a r k l e y S o u n d , 4 8 ° 4 8 . 8 ' N , 1 2 5 ° 1 0 . 6 » « , UEC 5 4 1 2 9 , 10 MAY 1 9 7 4 ; B o t a n i c a l B e a c h , 4 8 ° 3 2 ' N , 1 2 4 ° 2 7 V H , UBC 5 4 8 8 4 , 14 MAY 1 9 7 6 . washingtpn: Haatch Point, 2 JUN 1 9 6 6 .  Mukkaw  Bay,  4 8 ° 20»N,  1 2 4 ° 4 0 ' H , UBC 2 4 8 8 2 ,  Or e ^ o n : I n d i a n B e a c h , C l a t s c p C o u n t y , 4 5 ° 5 6 » N , 1 2 3 ° 5 9 « « , UBC 4 7 3 2 8 , 14 HAY 1 9 7 2 ; C h a p m a n P o i n t , C l a t s o p C o u n t y , 4 5 ° 5 5 ' N , 1 2 3 ° 58'W, UBC 4 7 5 8 4 , 12 J U N 1 9 7 2 ; S h o r t S a n d B e a c h , 4 5 ° 4 5 * N , 1 2 3 ° 58'W, UBC 2 4 8 8 3 , 6 J U N 1 9 6 6 . California: P i l l a r P o i n t , S a n M a t e o C o u n t y , 3 7 ° 3 0 * N , 1 2 2 ° 3 0 ' 8 , RS 4 7 4 , 10 J U L 1 9 6 8 ; P o i n t J o e , M o n t e r e y P e n i n s u l a , 3 6 ° 3 6 . 6 » N , 1 2 1 ° 57.4'W, G J B 2 6 7 2 , 18 J U N 1 9 3 9 ; UBC 5 4 8 1 4 , 10 J U L 1 9 7 6 ; P e b b l e B e a c h , M o n t e r e y P e n i n s u l a , 3 6 ° 3 4 * N , 1 2 1 ° 5 7 ' I , GMS 1 2 4 6 1 , 7 MAY 1 9 7 3 ; M i s s i o n P o i n t , C a r m e l , 3 6 ° 3 2 . 5 ' N , 1 2 1 ° 5 6 * W , GMS 4 7 6 3 , 16 MAR 1 S 6 6 ; E s t e r o P o i n t , S a n L u i s O b i s p o C o u n t y , 3 5 ° 2 8 » N , 1 2 0 ° 5 8 » W , AHF 8 0 4 3 7 , 6 AUG 1 9 7 4 .  237  Mexico : Punta Banda, B a j a C a l i f o r n i a , 77063, 25 JUL 1971. Spermatangial  and  3 1 ° 44»N, 1 1 6 ° 44*8,  AHP  carposporangial  Alaska: Bingham I s l a n d , 6 0 ° 01*N, 144° 2 3 » W , OBC 25459, 4 JUL 1966; K l o k a c h e f I s l a n d , 57° 25»N, 135° 53*8, OBC 23397, 30 JUN 1965; L o r a n S t a t i o n , B i o r k a I s l a n d , 5 6 ° 51* N, 1 3 5 ° 32*8, OBC 10291, 11 JUL 1960. British  Columbia:  T a s u N a r r o w s , Tasu Sound, Queen C h a r l o t t e I s l a n d s , 52° 4 4 , 5 » N , 132° 06«W, UBC 53651, 17 AUG 1970; T r e e I s l e t s , 50* 58.9*N, 127° 42.6*W, UBC 54888, 24 JUN 1975; Cape S c o t t , 50° 47*N, 128° 25*8, UBC 35742, 11 AUG 1968; S o l a n d e r I s l a n d , 50° 06. 5*N, 1 2 7 ° 56.3*8, UBC 53836, 25 JON 1975; Lynne Bock o f f C a u t i o u s P o i n t , 5 0 ° 04.3*N, 127° 3 3 . 4 » W , BBC 36036, 16 AUG 1968; D i a n a I s l a n d , B a r k l e y Sound, 4 8 ° 50.1»N, 125° 11.1*8, OBC 54821, 13 MAI 1976; UBC 54820, 16 JUN 1975; UBC 54136, 17 JUN 1975; UEC 54130, 19 JUL 1974; UBC 54137, 8 AUG 1975; UBC 54122, 14 SEP 1974; UBC 54132, 17 OCT 1974; Cape B e a l e , 48° 47*N, 125° 13*8, UBC 54891, 28 JUL 1976; B o t a n i c a l B e a c h , 4 8 ° 32'N, 124° 27»W, UBC 54133, 11 JUL 1975. Washington: Waadah I s l a n d ,  4 8 ° 23*N, 1 2 4 ° 36*W, UBC 8767,  15 AUG  1958.  Oregon: I n d i a n Beach, C l a t s o p C o u n t y , 4 5 ° 56*N, 1 2 3 ° 5 9*8, UBC 49873, 1 JUL 1973; J o c k e y Cap Bock, C l a t s o p C o u n t y , 4 5 ° 51*N, 123° 58*8, UBC 48333, 8 JUL 1972. California: T r i n i d a d H a r b o r , Humboldt C o u n t y , 4 1 ° 03* N, 124° 09'8, CSUH 07353, 12 APB 1972; Bodega Head, Sonoma C o u n t y , 3 8 ° 18*N, 123° 03*W, UBC 20010, 21 JON 1963; T o m a l e s Bay, M a r i n C o u n t y , 38° 14«N, 1 2 2 ° 55'W, AH J 55944, JON 1915; An"o Nuevo I s l a n d , 37° 06*N, 122° 20*8, Hansen 1631 i n OCSC { f i l e d u n d e r L a j d n a r j j t s i n c l a i r i i ) , 8 AOG 1972; D a v e n p o r t , 37° 00*N, 1 2 2 ° 11*W, GflS 2083, 6 J 0 L 1965; P o i n t J o e , M o n t e r e y P e n i n s u l a , 36° 3 6 . 6 » N , 121° 57.4*8, OBC 54818 and OBC 54887, 10 JOL 1976; P e s c a d e r o P o i n t , 36° 33.8*N, 121° 57*8, OBC 54892, 12 JOL 1976; M i s s i o n P o i n t , C a r m e l , 3 6 ° 3 2 . 5 » N , 121° 56'W, OBC 5 4 8 1 5 , 13 JOL 1S76.  238  Porphvra  nereocystis  Alaska: S t a r a y a Bay, U n a l a s k a I s l a n d , A l e u t i a n I s l a n d s , 5 3 ° 37*N, 166° 32* W, UBC 2677 6, 16 J U L 1966; K u p r e a n o f P o i n t , K o d i a k I s l a n d , 5 7 ° 55* N, 1 5 3 ° 05*H, UBC 26949, 20 JUL 1966; B o u s s c l e Head, 5 8 ° 23*N, 136° 5 5 * 1 , UBC 21199, 29 JUN 1965. British  Columbia:  L a n g a r a I s l a n d , Queen C h a r l o t t e I s l a n d s , 5 4 ° 13.54*N, 132° 58.6*W, UBC 56677, 29 MAH 1S76; M a s s e t , Graham I s l a n d , Queen C h a r l o t t e I s l a n d s , 5 4 ° 00 »N, 1 3 2 ° 09'8, UBC 50229, 10 MAS 1973; O f f N. W. c o a s t o f C a l v e r t I s l a n d , 5 1 ° 4 2 » N , 128° 05«W, UBC 42946, 13 APB 1970; T r e e I s l e t s , 50° 58.9'N, 1 2 7 ° 42.6'W, UBC 54105, 24 JUN 1975; Hope I s l a n d , 5 0 ° 54*N, 127° 57 »W, OBC 57182, 15 APB 1975; A m p h i t r i t e P o i n t , V a n c o u v e r I s l a n d , 48° 54'N, 1 2 5 ° 33'W, UBC 39820, 8 MAY 1969; A g u i l a r P o i n t , B a r k l e y Sound, 4 8 ° 50.4»N, 1 2 5 ° 08.3»W, UBC 57183 a n d 57184, 2 APB 1977; C a b l e Beach, B a r k l e y Sound, 4 8 ° 49.6'N, 1 2 5 ° 0 9 . 1 « H , UBC 57181, 8 AUG 1974; Second B e a c h , B a r k l e y S o u n d , 4 8 ° 48.9*N, 125° 10'H, UBC 57186, 9 MAY 1974; D i a n a I s l a n d , B a r k l e y Sound, 48° 5 0 . 1 » N , 1 2 5 ° 11.1 'SI, UBC 57185, 12 DEC 1976, UBC 57186 and 57189, 23 JAN 1977, UBC 57187 and 57190, 23 PEB 1975; L e a c h I s l e t , B a r k l e y Sound, 48° 49.8*N, 1 2 5 ° 14.4'w, UBC 57191, 2 APB 1977; B o t a n i c a l B e a c h , 4 8 ° 32 *N, 1 2 4 ° 27'W, UBC 50652, 25 APB 1974, UEC 57193, 11 J U L 1975; E s g u i m a l t , V a n c o u v e r I s l a n d , 48° 28'N, 1 2 3 ° 26'H, UC 96517, no d a t e . British  Columbia - S t r a i t  o f Georgia  Maude I s l a n d , 49° 16*N, 124° 05* W, UBC 11583, 15 AUG 1959; T u g b o a t I s l a n d , 4 9 ° 0 9 « N , 1 2 3 ° 41 »W, UBC 56603, 29 DEC 1976; G o w l l a n d P o i n t , S o u t h P e n d e r I s l a n d , 4 8 ° 44.0* N, 1 2 3 ° 11.2*8, UBC 57194, 4 SEP 1977; S i d n e y , V a n c o u v e r I s l a n d , 4 8 ° 39*N, 123° 2 4 » B , UBC 1434, 26 SEP 1917. Washington: F r i d a y H a r b o r , San J u a n I s l a n d , 48° 32*N, 123° 00*W, OBC 57195, 27 OCT 1S73; Whidbey I s l a n d , 4 8 ° 1 3 » N , 1 2 2 ° 46'W, UEC 1244, no d a t e . California: P e b b l e B e a c h , Monterey P e n i n s u l a , 57196, 4 NOV 1975.  3 6 ° 34*N, 121° 57*W, UBC  239  Porphyra  British  thuretii  Columbia:  S h i e l d s B a y , R e n n e l l Sound, Queen C h a r l o t t e I s l a n d s , 530 22.65*8, 132° 30.7'W, UBC 55302, 30 MAR 1976; V o l c a n i c C o v e , K y u g u o t C h a n n e l , V a n c o u v e r I s l a n d , 4 9 ° 58.7*8, 127° 13.9*8, UBC 54446, 15 HAY 1975; D i a n a I s l a n d , B a r k l e y Sound, 48° 50.1*N, 125° 11.1'W, UBC 57203, 23 JAN 1977; UBC 52019, 23 FEB 1975; UBC 57202, 29 APR 1975; L e a c h I s l e t , B a r k l e y Sound, 4 8 ° 49.8* N, 125° 14.4*8, UBC 57200 a n d 57201, 2 APR 1977. California: P a c i f i c G r o v e , M o n t e r e y C o u n t y , 36° 3 7 . 6 » N , 1 2 1 ° 55* K, UC 95610, 23 APR 1897; UC 95598 and 95596, JUN 1901; UC 95612, no d a t e ; P e b b l e Beach, H o n t e r e y P e n i n s u l a , 3 6 ° 34*N, 1 2 1 ° 57*8, GHS 1406, 24 JUN 1943; C a r m e l Bay 36° 33*N, 1 2 1 ° 56*8, 0C 791973 and 791970, 29 MAY 1900; GMS 6467, 23 JUN 1939.  

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