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Some effects of ultraviolet radiation on fish Bell, Gertrude Mary 1949

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(in fat  ftf sc  SOME EFFECTS OF ULTRAVIOLET RADIATION ON  FISH  by  G e r t r u d e Mary  A Thesis the  Bell  Submitted i n P a r t i a l  Fulfilment  Requirements f o r the Degree o f M A S T E R in  OF  A R T S  the Department of ZOOLOGY  The U n i v e r s i t y  of B r i t i s h Columbia  APRIL,  1949  of  TABLE OP  CONTENTS  INTRODUCTION  v  HISTORICAL REVIEW  1  Ultraviolet  light  .  1  Ultraviolet  i n sunlight  •  1  Some b i o l o g i c a l  effects  of u l t r a v i o l e t  radiation  •  2  E a r l y work u s i n g u l t r a v i o l e t  4  Effect  on human b e i n g s  6  Effect  on eggs  8  Effect  o f X - r a d i a t i o n on f i s h  8  METHODS AND  MATERIALS  Sources  10  of U l t r a v i o l e t  Standardization  o f lamps  10 10  Experimental animals  12  Methods  13  Histological  studies  C o m p a r i s o n o f lamps  14 and s u n l i g h t  RESULTS  15 19  Preliminary  experiments  19  Chum s a l m o n eggs  21  Sockeye  s a l m o n e ggs  24  Sockeye  salmon a l e v i n s  30  Histological  studies  34  (A)  Skin  34  (B)  Other t i s s u e s  40  ii  DISCUSSION  41  Ultraviolet  and r a t e  Mortalities  i n eggs a n d a l e v i n s  Effects  of u l t r a v i o l e t  on f i s h  41 41  skin  .... 44  (A)  Preliminary  (B)  A l e v i n sockeye salmon e x p e r i m e n t s . . . . . . . . . . . . . . . . . . . 46  Effects SUMMARY  of hatching  .  e x p e r i m e n t s ....... 44  of ultraviolet  o n o t h e r t i s s u e s . 49 ..  51  ACKNOWLEDGMENTS  52  LITERATURE CITED  53  iii L I S T OF  ILLUSTRATIONS  TABLES Table  - A v e r a g e h o u r s o f a c t u a l s u n s h i n e and o f e r y t h e m a l u l t r a v i o l e t energy as measured at Cleveland, Ohio.  I  Table I I  - T o t a l number of E - v i t o n h o u r s upon a l e v l n s o c k e y e s a l m o n .  Table I I I  - Percentage m o r t a l i t i e s  o f chum s a l m o n  T a b l e IV  - Percentage m o r t a l i t i e s eggs.  of sockeye  incident eggs,  salmon  Table  V  - R e m o v a l s , d a i l y m o r t a l i t i e s and c u m u l a t i v e percentage mortality of a l e v i n sockeye salmon.  Table  VI  - C a l c u l a t e d t h e o r e t i c a l deaths of c o n t r o l and i r r a d i a t e d a l e v i n s o c k e y e s a l m o n .  FIGURES Figure  1  - Some b i o l o g i c a l e f f e c t s radiation.  of u l t r a v i o l e t  Figure  2  - Graphs salmon  of percentage mortality eggs.  o f chum  Figure  3  - Graphs salmon  of percentage m o r t a l i t y eggs.  o f sockeye  Figure k  - Sockeye s a l m o n a l e v i n s irradiation.  i+6 d a y s  Figure 5  -  Figure  6  - S k i n o f a l e v i n sockeye salmon 3 d a y s a f t e r i r r a d i a t i o n o f tac.  Figure  6a  - Daily cumulative m o r t a l i t i e s salmon a l e v i n s .  Figure  7  - S k i n o f c o n t r o l s o c k e y e salmon 7 d a y s a f t e r experiment began.  Figure  8  - Skin of f i s h o f 2x.  C o n t r o l sockeye salmon a l e v i n s a f t e r the experiment began.  k days  after k6  days  of sockeye  a f t e r an i r r a d i a t i o n  iv  Pig.  9  - S k i n of f i s h o f 8x.  4 days a f t e r  an  irradiation  Pig.  10 - S k i n o f f i s h o f 0.5x.  7 days a f t e r  an  irradiation  Pig.  11 - S k i n o f f i s h o f 2x.  7 days a f t e r  an  irradiation  Pig.  12  13 days  - Skin of f i s h o f 0.5x.  after  a dosage  V  SOME E F F E C T S OF ULTRAVIOLET RADIATION ON  FISH  INTRODUCTION The e f f e c t s o f u l t r a v i o l e t h a v e studied Little  on man  been  and on t h e l o w e r f o r m s s u c h as  o r no work h a s b e e n done on t h e l o w e r  particularly  aquatic  forms.  Such s t u d i e s  extensively bacteria.  vertebrates,  will  i n t e r e s t i n g f r o m t h e c o m p a r a t i v e a n g l e and may practical  significance, since  young h a t c h e r y f i s h may  prove be o f some  i t has been s u g g e s t e d  be a d v e r s e l y  affected  by  sunlight.  The w a v e l e n g t h s u s e d a r e a p p r o x i m a t e l y t h e same as found i n the u l t r a v i o l e t of  have been i n v e s t i g a t e d  (1)  as  l e t h a l e f f e c t s on a d u l t auratus), and  those  sunlight.  I n t h i s s t u d y the e f f e c t s o f t h e n e a r violet  that  ultra-  follows: goldfish  (Carassius  coho s a l m o n f r y ( O n y c o r h y n c h u s k l s u t c h ) ,  d e v e l o p i n g eggs o f chum s a l m o n ( O n y c o r -  h y n c h u s k e t a ) and s o c k e y e s a l m o n  (Onycorhynchus  nerka). (2)  h i s t o l o g i c a l c h a n g e s i n t h e s k i n and tissues  (3)  of sockeye salmon  underlying  alevins.  colour  changes p r o d u c e d i n f i s h  violet  radiations.  s k i n by  ultra-  1.  HISTORICAL REVIEW  Ultraviolet  light  Ultraviolet three  regions,  ultraviolet. finds is  little  light  i s commonly d i v i d e d , i n t o  the near, middle  and f a r o r extreme  The e x t r e m e u l t r a v i o l e t biologic  below 2000 A  and t h e r a p e u t i c use s i n c e i t  a b s o r b e d by most s u b s t a n c e s i n c l u d i n g a i r .  middle u l t r a v i o l e t in biologic mitted  from  2000 A t o 3000 A i s i m p o r t a n t  and t h e r a p e u t i c w o r k .  I t i s not t r a n s -  by g l a s s b u t i s t r a n s m i t t e d  glasses  free of iron.  extends  into this  2900 A.  Ordinary  by q u a r t z  The s o l a r s p e c t r u m  region,  and  fluorescence  from  3000 A t o a b o u t  b e t w e e n 3900 A and  3500 A, b u t t h e t r a n s p a r e n c y r a p i d l y d i m i n i s h e s most g l a s s e s  of ordinary  thickness  at  The v i s i b l e  spectrum  o f about  3900 A, d e p e n d i n g  (Luckiesh,  about  and p h o t o g r a p h i c a c t i o n .  glass i s quite transparent  3100 A.  special  barely  the lower l i m i t being  The n e a r u l t r a v i o l e t  4000 A p r o d u c e s  The  are f a i r l y  until  opaque  begins at a  wavelength  on t h e s e n s i t i v i t y  o f t h e eye  1946).  Ultraviolet  i n sunlight  The u l t r a v i o l e t  components o f s u n l i g h t  t h a t r e g i o n o f the spectrum imately  2850 A, t h a t  visible  spectrum  from  about  i s , from t h e l o w e r  i n the blue  include  4000 A t o a p p r o x regions  t o the lowest  o f the  wavelengths  t r a n s m i t t e d by t h e transmitted of and  day,  e a r t h ' s atmosphere.  to the e a r t h v a r i e s  t h e l a t i t u d e , the  other f a c t o r s ,  2950 A. solar  Laurens  energy  the  w i t h the  altitude,  average  The  season,  r a n g i n g from  sunlight  u  after  traversing  (3200 A)  and  the  constitute  a most v a r i a b l e  Some b i o l o g i c a l The  effects  lamps were f o u n d  radiations of  egg  are absorbed  skin  t o form  part  visible  of t o t a l  o f i t " (Blum,  1945).  radiation  of the v a r i o u s  spectral  Germicidal  i f the  by b a c t e r i a .  The  energy these  The c o a g u l a t i o n  activation  V i t a m i n D was 3100  A.  as  shorter  t h a n 0.2$  to d i f f e r markedly.  t o t h e most e f f e c t i v e  found  of  have  ergosterol  to h a v e a p e a k  This corresponds  fairly  l i n e s which produce o f untanned  human  eryskin  1). Other  the  less  of u l t r a v i o l e t  thema o r t h e r e d d e n i n g e f f e c t (Pig.  i n the  the p r o d u c t i o n of c o n j u n c t i v i t i s  2500 A.  between 2500 A and closely  to  c o n c e n t r a t e d a t 2500 A t o 2600 A s i n c e  a maximum n e a r  time  e a r t h ' s atmosphere  t o be most e f f e c t i v e  albumen and  i n the  2850 A  "Wavelengths  biological effects  r e g i o n s have been f o u n d  e m i t t e d was  the  (1928) g i v e s t h e d i s t r i b u t i o n o f the  59 p e r c e n t i n t h e i n f r a r e d .  t h a n 0.32  limit  atmospheric c o n d i t i o n s  1 p e r c e n t i n t h e u l t r a v i o l e t , 40 p e r c e n t and  lowest  lines  i n the  spectrum  same r e s u l t s , b u t w i t h l e s s  a b s o r p t i o n o f the  particular  are found  efficiency,  substance  to  since  concerned  produce the  falls off.  Pig.  1 - G r a p h s h o w i n g t h e r e l a t i v e e f f e c t s o f u l t r a v i o l e t r a d i a t i o n i n the n e a r m i d d l e u l t r a v i o l e t (redrawn f r o m L u c k i e s h , 1 9 4 6 ) . Arrows i n d i c a t e wavelengths used i n t h e s e e x p e r i m e n t s .  and  4.  The  a c t i v a t i o n of e r g o s t e r o l  of erythema f a l l w i t h i n t h e violet  transmitted  germicidal  by  u l t r a v i o l e t has hormones.  the  and  are  now  reaction.  enzyme i s p r o b a b l y due  o f the  p r o t e i n m o i t y of the. enzyme.  not  by  (1917) o b s e r v e d t h a t  p a r a m e c i a by  s i m i l a r to  Harris  and  (1917) f o u n d  T h e s e s u b s t a n c e s may  which render l i v i n g of u l t r a v i o l e t  was  when a b s o r b e d may  of the  coagulation  heat.  that  tyrosine  bands i n the as  susceptible  kills  protoplasm  The  and  ultra-  optical sensitizers to  the  (1922) s u g g e s t e d t h a t  the  toxic  a photolytic process. cause s e v e r a l  effect  The  action  of  radiations  effects*  (1) L o n g e r r a d i a t i o n s w i t h r e l a t i v e l y of  denaturation  light.  Clark ultraviolet  cells  act  inacti-  ultraviolet  enzymes.  absorption  and  have  The  t h a t p r o d u c e d by  Hoyt  phenylalanine exhibited violet.  that  which  to the  coagulation  i n a c t i v a t i o n o f the  p r o d u c e d was  known t o  group a p r o t e i n  of  and  most e f f e c t i v e  (1916) f o u n d  vation  b a c t e r i a and  ultra-  sunlight.  Neil  s p e c i f i c i t y o f the  Burge  the  power t o i n a c t i v a t e enzymes  to t h e i r p r o s t h e t i c  the  production  ultraviolet  These substance©  governs the  of  the  of  s u n l i g h t , w h i l e the  Burge, F i s c h e r  attached  lower l i m i t s  area i s below that  E a r l y work u s i n g  and  energy, i . e . , i n f r a r e d , v i s i b l e  small and  amounts near  5.  ultraviolet,  when absorbed, may c a u s e  molecular motion, producing a r i s e  increased  i n temper-  ature. (2)  When r a d i a t i o n o f s u f f i c i e n t fluorescent electrons cules,  substance  energy f a l l s  on a  I t has been supposed  are separated  from t h e i r p a r e n t  c r e a t i n g an u n s t a b l e  condition.  that mole-  Recom-  b i n a t i o n o f the e l e c t r o n o c c u r s w i t h t h e emission of l i g h t . the an  visible  The l o w e s t w a v e l e n g t h s o f  and near u l t r a v i o l e t  produce  such  effect.  (3) W i t h t h e s h o r t u l t r a v i o l e t  region,  X - r a y s and  radioactive p a r t i c l e s a photochemical e f f e c t Is produced. by  The e l e c t r o n s  the r a d i a t i o n s that  t h e i r parent•molecule w i t h some o t h e r Clark  living  cells  they a r e separated  a n d a r e f r e e t o combine  energy s h o r t e r  thereby l e a d i n g to a chemical  Light  t h a n 3000 A a c t e d  presence  reaction.  i n the same way i n t h e  o f a s e n s i t i z e r , w h i c h so a f f e c t e d the  conditions threshold  t h a n 3000 A  by i o n i z i n g the p h o t o e l e c t r i c a l  constituents longer  from  atom o r g r o u p o f a t o m s .  (1922) b e l i e v e d t h a t  affected  a r e so a c t i v a t e d  o f these constituents  that  s h i f t e d i n t o the v i s i b l e .  surface  their photoelectric She b e l i e v e d  that  m e l a n i n pigment formed i n t h e s k i n by s u n l i g h t acted a s e n s i t i z e r , s h i f t i n g the p h o t o e l e c t r i c p o i n t  as  Into the  6. near  ultraviolet.  Effect  on human b e i n g s In  s m a l l doses resulting while  human b e i n g s  i t has been demonstrated  of ultraviolet  c a n produce  increase i n pigmentation  s t r o n g e r doses  a c t e d upon.  c a n cause  that  erythema w i t h a  i n s k i n and h a i r ,  n e c r o s i s of the t i s s u e  Wounds h a v e b e e n shown t o h e a l more  rapidly,  s k i n f o r m s o f l u p u s h a v e b e e n a i d e d o r c l e a r e d up, and the  antirachitic  effect  Irradiations  i s w e l l known  (Laurens,  of experimental animals  beings w i t h a r t i f i c i a l  1933).  and human  s o u r c e s h a v e shown s l i g h t and hemoglobin content  effect  on r e d b l o o d c e l l  count  of blood  when t h e s e l e v e l s  a r e n o r m a l a t t h e b e g i n n i n g o f the  i n v e s t i g a t i o n s . Anemic c o n d i t i o n s h a v e b e e n a i d e d , t h e number o f e r y t h r o c y t e s , l y m p h o c y t e s  and e s p e c i a l l y  p l a t e l e t s have been i n c r e a s e d s u b s t a n t i a l l y . has  n o t been a i d e d , b a s a l m e t a b o l i s m  have n o t been s u b s t a n t i a l l y light  (Mayerson,  a f f e c t e d by u l t r a v i o l e t  light  has been c o r r e l a t e d w i t h t h e  production of s k i n cancers, both  pointed in  out that  and t h e t h y r o i d  1935).  Ultraviolet  i n human b e i n g s  carcinomas  and e x p e r i m e n t a l a n i m a l s .  more common i n o u t d o o r i n those  r e c e i v i n g more  (1945)  principally  and t h a t  workers than indoor  areas  and sarcomas, Blum  cancer of the s k i n occurs  p a r t s o f t h e body exposed t o s u n l i g h t  especially  Growth  i t is  workers,  sunlight.  7.  Cancer of the by  s k i n has  been p r o d u c e d  exposure to u l t r a v i o l e t  therapy, b e l i e v e d  t h a t pigment  organism  excessive  p r o t e c t i o n which i t a f f o r d s  the  important  radiation, arising  response t o  as  an  i t d o e s , as  inflamed,  while  and  thus p r o t e c t e d , Since  1928  skin to ultraviolet Laurens  showed an the  cutaneous r e s i s t a n c e the  to the  intense  adaption  ultraviolet  the  factor that  of p e n e t r a t i o n  the  radiant  energy.  t o be  a surface  lengths  l e s s than  whereas a t 2500 A  2800 A a r e and  more r a d i a t i o n r e a c h i n g corium.  the  living  that  the  increased  d e t e r m i n e d the The one  degree  reaction  the  horny is  the  absorbed  skin, while  layers,  greater,  s t r a t u m M a l p i g h i i and  l a y e r s of  of  s i n c e wave-  the p e n e t r a t i o n  W a v e l e n g t h s b e l o w 2500 A a r e  they reach the  that  involved  a b s o r b e d by  3000 A,  o f human Subsequently  (1945) b e l i e v e d  l a y e r o f p i g m e n t and  human s k i n I s b e l i e v e d  glass,  reaction."  c o r n i f i c a t I o n was of  on  c o v e r e d by  r a d i a t i o n have changed. Blum  demonstrated  t h e p i g m e n t e d a r e a s were  views of the  and  physio-  showing t h a t  those p r e v i o u s l y  (1933, 1935)  skin outside  arm  pigment against  a  Pinsen  an (1928)  of  organism  overstimulation.  exposures of the  not  Laurens  functions  p r o t e c t i v e a c t i o n o f p i g m e n t by  successive  ultraviolet protected  radiation.  is  logical  of  formation  that,  excessive  o f the  field  stated the  "One  rats  radiation.  E a r l i e r workers i n the  against  i n m i c e and  the before  wavelengths  8.  longer  t h a n 3000 A h a v e b e e n f o u n d  considerable  Effect  on  distances  patterns.  germinal  i n the  shown t o be  being  tail  the  Pundulus  produced early  cleavage  i n t h e h e a d , e y e s and  irradiations  r i n g was  cleavage  eggs o f  with u l t r a v i o l e t  Irradiations during  regions, while  abnormalities be  (1925) f o u n d t h a t  caused a b n o r m a l i t i e s  when t h e  1933).  b e e n shown t o a l t e r  irradiated  abnormalities.  may  has  Hinricks  heterolltlcus  cardial  to  eggs Ultraviolet  stages  (Laurens,  to p e n e t r a t e  d u r i n g the  laid  region.  time  down p r o d u c e d  These  consistent with  peri-  abnormalities  the metabolic  gradients,  t h e most a c t i v e a r e a m e t a b o l i c a l l y b e i n g the  most a f f e c t e d  when an  she  out, but  inhibiting  the may  abnormalities be  pressure, chemical  Effect  i n f l u e n c e Is a c t i v e . were n o t  p r o d u c e d by  a number o f  temperature,  increased  environment, X-rays,  o f X - r a d i a t i o n on Bonham e t a l  decreased  the  irradiation  cell  f a c t o r s such  as  alteration  that  of  X-radiations  c o u n t s 2 t o 3 weeks  irradiation  were r e d u c e d s i g n i f i c a n t l y used.  ultraviolet,  fish  cell  count i n the  in fingerling  salmon, Onycorhynchus t s c h a w y t s c h a .  which they  pointed  etc.  the hemopoietic  1 t o 2 weeks a f t e r  for  salinity,  (1948) f o u n d  red blood  and  specific  As  kidney  Chinook  W e i g h t and  i n the h i g h e r  after  length  irradiations  9.  Welander Influenced  e t a l (1948) showed t h a t  the growth  rate,  development  and  o t h e r o r g a n s o f embryo a n d l a r v a l  The  l a r g e r doses  which  eyes a n d  Subcutaneous  b e l l y w a l l below  the d e v e l o p -  system,  fins,  system a p p e a r e d  after  appeared  e x t r a v a s a t i o n was  the l a t e r a l  ment o f t h e y o l k s a c .  red  line  noted along  n e a r the a t t a c h -  A t t h e same t i m e w h i t e  on t h e y o l k  sac.  After  b l o o d seemed t o be c i r c u l a t i n g Histological  skeletal  revealed to  in  in  integumentary  .Studies cell  o f t h e k i d n e y was  o f the pronephros  counts w e l l below irradiation,  that  severely  of irradiated  of dose.  fish  o f the c o n t r o l s  a f t e r which  up  recovery took  R e d u c t i o n of the p e r c e n t a g e o f m i t o t i c  t h e h e m o p o i e t i c t i s s u e was  amount  o r no  systems.  23 d a y s a f t e r  place.  little  i n the normal c h a n n e l s .  studies revealed l i t t l e  Hemopoietic t i s s u e damaged.  flocculent  5 weeks  damage i n t h e n e r v o u s , d i g e s t i v e , m u s c u l a r , or  salmon.  growth.  33 d a y s .  spots  chinook  the v a s c u l a r  Damage t o t h e v a s c u l a r  the  o f t h e eye  they used a r r e s t e d  ment o f c u t a n e o u s p i g m e n t ,  X-rays  cells  i n p r o p o r t i o n to the  G l o m e r u l i were i n h i b i t e d  i n development  t h e m e s o n e p h r o s , b u t i n t e r r e n a l b o d i e s were n o t  damaged  as e x t e n s i v e l y  as h e m o p o i e t i c t i s s u e .  and s p l e e n were damaged h e a v i l y ,  Gonads  i n both cases the c e l l s  showed p y c n o s i s a n d s u b s e q u e n t d e g e n e r a t i o n .  10.  METHODS AND Sources of  ultraviolet  Preliminary cold  quartz-mercury  Products was  MATERIALS .  M i n e r a l i g h t lamp  I n c . , Los  operated  dominantly  e x p e r i m e n t s were p e r f o r m e d w i t h  Angeles,  w i t h the  (Ultra-violet  California).  filter  attached  and  at the mercury resonance l i n e ,  This  lamp  emits  pre-  2537  T h e s e p r e l i m i n a r y e x p e r i m e n t s were c a r r i e d t h e method o f dosage- s t a n d a r d i z a t i o n h a d The  b u l k of the  w i t h 275  watt G e n e r a l  on  volt  a  a 110 special  "The  AC  type  circuit.  The  lamp t a k e s  One  with  reflector  commonest  of f a u l t s  The  the  incorThe  intensity  (Weitz,  in  1946).  biological  total  energy  some n o n - s e l e c t i v e m e t h o d .  s h o u l d be m e a s u r e d i n r e l a t i v e by  discharge  type b u l b . light  A.  lamps  s t a n d a r d i z a t i o n of dosage.  thermopile,  i s of  resistance ballast  5 minutes f o r r e s t a r t i n g "  of the  a s c e r t a i n e d by  operating  a mercury  r e s e a r c h u s i n g u l t r a v i o l e t r a d i a t i o n s i s the  be  out  down t o 2800  about 3 minutes t o r e a c h f u l l  S t a n d a r d i z a t i o n of  developed.  g l a s s on t h e b u l b  a tungsten-filament  approximately  been  before  sunlamps,  transmitting ultraviolet  p o r a t e d w i t h i n an u l t r a v i o l e t  and  t y p e RS  lamp i s a s e l f - c o n t a i n e d u n i t  element and  out  A.  e x p e r i m e n t s were c a r r i e d  Electric  a  deflection  or  lack of should This  a b s o l u t e u n i t s by  of a galvanometer,  or  a by  11.  some p h o t o c h e m i c a l method w h i c h r e q u i r e s amount  o f energy  t o produce  a visible  a definite  change I n a  property of the s o l u t i o n . The this  study.  of h y d r i o d i c Influence give  method o f P o h l e I t i s based  (1926) h a s b e e n u s e d i n  on the f a c t t h a t  a c i d i n water frees  of intense  light.  a solution  i o d i n e u n d e r the  This  free  iodine  a blue c o l o r a t i o n with the starch t e s t .  will Details  o f t h e method u s e d a r e : (1)  The lamp was a l l o w e d t o warm u p 15 t o 20 minutes,  (2)  A large  or u n t i l  t h e o u t p u t was c o n s t a n t .  amount o f 5.3$ H2SO4 was made u p  sufficient  for a l l tests.  (3) N / l O sodium t h i o s u l f a t e was p r e p a r e d i n s u f f i cient  quantity  f o r a l l t e s t s by a d d i n g  24.83  gms. o f Na2Sg O3. 5E"20 t o 1000 gms. o f w a t e r . The  N/400 s o l u t i o n u s e d f o r e a c h t e s t was  made up by a d d i n g 1 c c . o f N / l O N a S 2 0 t o 2  39 c c . o f d i s t i l l e d  water.  (4) A 1% s o l u t i o n o f s o l u b l e  s t a r c h was p r e p a r e d  at the b e g i n n i n g o f each experiment s o l v i n g 1 gm. o f s o l u b l e cold d i s t i l l e d  starch  by d i s -  i n5 cc.  of  w a t e r a n d a d d i n g t h i s t o 95 c c  of b o i l i n g d i s t i l l e d (5) The p o t a s s i u m  3  iodide  water. was p r e p a r e d  freshlyf o r  e a c h t e s t by d i s s o l v i n g 250 mg. o f K I i n 25 c  12. of d i s t i l l e d  water.  (6) I n a 100 m l . b e a k e r o f 5 cm. u p p e r d i a m e t e r c c . o f K I p l u s 25 c c . o f 5,3%  H S0 2  plus  4  5  25 drops  o f 1% s t a r c h s o l u t i o n and 1 c c . o f N/400 N a S 2 0 2  were p l a c e d u n d e r t h e lamp. and  Blue  " c l o u d s " formed  were s t i r r e d w i t h a g l a s s r o d .  r e q u i r e d f o r t h e whole  E-viton,  incident  centimeter  p e r i o d was  the time  o f erythemal  f l u x or 1  upon a p r o j e c t e d a r e a  (Luckiesh, 1946).  r u n were c a l i b r a t e d  Tests  T h i s time  1 dos;e and i s e q u i v a l e n t t o 1 P i n s e n . 1  P i n s e n i s 10 m i c r o w a t t s  called  The t i m e  s o l u t i o n t o t u r n b l u e was  m e a s u r e d by a s t o p w a t c h . called  All  of 1  experiments  i n terms of t h i s  c o l o u r change,  f o r the s o l u t i o n t o t u r n b l u e  being  "1 d o s e " o r "1 x " . such  a s t h e above a r e v a l u a b l e when u s i n g a  mercury a r c s i n c e t h e y g i v e an i d e a of the i n t e n s i t y with  age o f t h e lamp, and a l s o  w h i c h c a n be  reproduced.  Experimental  animals  Stouffville,  when  decrease  g i v e a s t a n d a r d i z e d dose  G o l d f i s h u s e d were o b t a i n e d f r o m in  a s u p p l y house  O n t a r i o a n d were 6 t o 8 cm. i n l e n g t h  irradiated. Chum  parents hatchery  3  s a l m o n eggs w e r e t h e p r o g e n y o f s e v e r a l  o b t a i n e d from  Sweltzer  creek,  on t h e a f t e r n o o n o f November  near  Cultus  28, 1948.  Lake The  13.  sockeye  s a l m o n eggs and  a l e v i n s were t h e p r o g e n y  several parents  o b t a i n e d a t Weaver c r e e k on  16,  eggs w e r e t a k e n  1948.  The  transferred troughs  immediately  a t the  to the  October  i n t h e ' u s u a l way experimental  part  and  hatchery  university.  Coho s a l m o n f i n g e r l i n g s were o b t a i n e d Capilano r i v e r  of  and  o f J u l y and  Nelson  c r e e k d u r i n g the  from  latter  e a r l y p a r t of August.  Methods The troughs  until  chum s a l m o n e g g s were c a r r i e d a s h o r t time before  the  i n the  irradiation,  t h e y were removed t o aluminum c o a t e d p a n s o f 22.5 cm.  The  pans were f i t t e d  w i t h an i n l e t  a l l o w i n g running water to a depth  were p l a c e d on t h e b o t t o m o f e a c h p a n  glass to  rods.  The  d i s t a n c e from  t h e b o t t o m o f t h e p a n s was  p l a t e was eggs and violet  p l a c e d over the  cm.  lamp, s e r v i n g t o f i l t e r  from  band i s a p p r o x i m a t e l y  The  A.  out  In this  a b o u t 3100  eggs layer  by means o f the  A heavy  manner  lamps glass  ultrathe  ultraviolet  A t o 2800 A.  equal t o the u l t r a v i o l e t  This  components  spectrum.  Water a l l o w e d t o c i r c u l a t e  through  of  the  a l l the  e g g s w e r e e x p o s e d t o a band o f  r a d i a t i o n extending  solar  the bottom  26.5  sq.  In a single  s q . cm.  when  outlet  c o n t r o l group between  r a y s b e l o w a b o u t 3100  experimental  o f the  the  and  o f 4 cm.  and were c o n f i n e d t o an a r e a o f 10  hatchery  the  pans  14. kept  the temperature  the experiments  had  c o n s t a n t w i t h i n 1° been c a r r i e d  ferred  t o an aluminum  counts  were made. The  eggs  out t h e  coated tub.  of the  experimental hatchery  sockeye  troughs.  i r r a d i a t e d under e x a c t l y the  t o 3°  C.  After  eggs were  Periodic  trans-  mortality  were m a i n t a i n e d  i n the  T h e s e eggs were  same c o n d i t i o n s as  the  chum s a l m o n eggs d e s c r i b e d a b o v e . The  a l e v i n sockeye,  on t h e  t r a n s f e r r e d from the t r o u g h s e v e r a l had  ceased  cm., The  days a f t e r h a t c h i n g  i n t o aluminum c o a t e d mesh b a s k e t s , 13  s u s p e n d e d i n the  trough t o a depth  f i s h were i r r a d i a t e d  there u n t i l was  o t h e r h a n d , were  completion  of 4 to 5  i n these baskets  of the  and  experiments.  cm.  The  lamp  the  a manner t h a t t h e d i s t a n c e f r o m  the  b o t t o m o f the lamp t o t h e b o t t o m o f t h e b a s k e t  was  26  cm.  The  lamp was  a l l o w e d t o warm up  before the i r r a d i a t i o n s  commenced.  e a c h g r o u p were t a k e n and  to f u l l  Daily  20  remained  p l a c e d s e c u r e l y i n a stand which straddled  trough i n such  x  intensity  samples  d a i l y m o r t a l i t i e s were  of  counted.  Histological studies Histological and  control  acetic acid to  fish fixed  i n Bouin's  fixative.  4 w a s h i n g s o f 10%  carbonate.  s t u d i e s were made o f t h e  The alcohol  . Dehydration  and  irradiated  p i c r i c acid-formol-  Bouin's  was  removed w i t h  saturated with c l e a r i n g was  3  lithium  done i n 3  changes  15. of d i o x a n and the manner.  f i s h were embedded i n wax  Saggittal  serial  s e c t i o n s were made a t 5  U n f o r t u n a t e l y t h e m i c r o t o m e u s e d was markedly  i n t h e t h i c k n e s s o f the  accurate c e l l the  b u t was the  less  Harris  1  sections  so  that  Removal  some damage t o the h e a d k i d n e y  The  h a e m o t o x y l i n and  of  region,  t h a n t h e damage c a u s e d by a l l o w i n g  yolk to remain.  u.  observed to vary  c o u n t s c o u l d n o t be made.  yolk caused  i n the u s u a l  some o f  s e c t i o n s were s t a i n e d i n e o s i n made up  i n the  prescribed  manner. Comparison  o f lamps and  sunlight  S i n c e t h e band experiments effective it  of u l t r a v i o l e t  c e n t e r e d about t h a t which  and  since  i s possible  the dosage  was  used  Is e r y t h e m a l l y  calibrated  t o make a r o u g h c o m p a r i s o n  ultraviolet  sunlight. The  Hltraviolet  of sunlight  erythemally effective units (Pinsens) Incident period  of t i m e .  p e r s q . cm. in  In Pinsens,  of the e n e r g i e s  u s e d i n t h e s e e x p e r i m e n t s w i t h t h o s e o f the in  i n these  exposure  o f 2/3  on t h e a v e r a g e u n t a n n e d  a minimum p e r c e p t i b l e Luckiesh  area f o r a  E-vitons  certain  o f an E - v i t o n human s k i n  In  hour  results  erythema.  (1946) h a s made a number o f m e a s u r e -  ments o f t h e u l t r a v i o l e t suburb  s u c h as t h e number o f  upon a p r o j e c t e d An  is calibrated  components o f s u n l i g h t  of C l e v e l a n d , Ohio.  A l t h o u g h the v a l u e s  in a which  16. he  gives  c o u l d be  (Table I ) are s l i g h t l y h i g h e r expected  comparison  i n Vancouver,  w i t h the  ultraviolet  than those  which  they a r e c o n v e n i e n t f o r used  i n these  experi-  ments .  Table  I -  Variation  actual  s u n s h i n e p e r month and  energy  i n daylight  for  ( a f t e r L u c k i e s h , 1946,  in a  of erythemal  p.  suburb  minutes,  71 98 167 221 298 305 350 292 243 178 103 66  1.8 2.5 6.3 9.8 15.0 16.7 16.7 13.5 9.8 5.0 1.9 1.0  were m e a s u r e d by  In A p r i l  the  time  Energy  Percent of t o t a l f o r year  51 72 182 280 430 476 476 386 277 143 55 30  a minimum p e r c e p t i b l e while  plane  Erythemal U l t r a v i o l e t  Seasonal variations  produce  ultraviolet  of C l e v e l a n d , O h i o  E - v i t o n hours p e r s q . cm.  (1946.) a t C l e v e l a n d .  of  57).  Average hours of sunshine  January February March April May June July August September Octobe r November December  hours  as m e a s u r e d on a h o r i z o n t a l  a s i x year p e r i o d  Month  of average  Luckiesh  required  to  e r y t h e m a a t n o o n was  a t n o o n i n June o n l y 17 m i n u t e s  50  were  necessary. The the sockeye  e r y t h e m a l v a l u e s o f the r a d i a t i o n s u s e d f o r s a l m o n a l e v i n s ware c a l c u l a t e d .  The  area  17.  o f t h e b a s k e t s u s e d was calibrated  260  s q . cm.  a t 1 P i n s e n i n 30 m i n u t e s .  F i n s e n hours x the s u r f a c e a r e a In E - v i t o n hours per sq. Table  II -  i n c i d e n t upon a l e v i n the  and t h e dosage  The  The  i n cm.  number o f the  dosage  cm. total  number o f E - v i t o n h o u r s  sockeye salmon  at the surface  of  water. C a l i b r a t i o n o f dose  T o t a l cLosage i n E - v i t o n h o u r s on an a r e a o f 260 s q . cm.  0.5x l.Ox 2.Ox .4.Ox 8.Ox The  of t h e w a t e r , r e c e i v e d the  65 130 260 520 1040  a c t u a l energy reaching  small f r a c t i o n  o f the I n c i d e n t s i n c e the s u r f a c e  r a d i a t i o n was  t o t a l surface  light  Bornstein,  area  surface  of each f i s h  absorption  which  of u l t r a v i o l e t  i n d i c a t e d t h a t f o r a d e p t h of 5  incident intensity  i s transmitted  cm. (Landolt-  1923).  Dissolved organic absorption  materials  of the u l t r a v i o l e t  ments o f V a n c o u v e r Department  e n e r g y on t h e  a  area.  i n water  94.5$ o f t h e  e a c h f i s h was  v e r y much s m a l l e r t h a n the  Measurements o f t h e  the  gives  was  and m i n e r a l s  increase  t o some e x t e n t .  Measure-  t a p w a t e r , made by D r . C r o o k e r o f the  of P h y s i c s ,  i n d i c a t e d that the values  f o r the  18.  absorption of ultraviolet differ  significantly  Bornstein  fish  from t h o s e  c a l c u l a t e d values  (Table  of the order  i n Landolt-  of energy i n c i d e n t upon  I I ) u s i n g t h e RS sunlamp were p r e s u m a b l y  which t h e y  summer s u n l i g h t .  could  receive  Thus the e f f e c t s  e x p e r i m e n t s a r e ones w h i c h hatchery  given  (1923). The  the  energy i n t a p water d i d not  conditions.  i f exposed t o  observed i n these  c o u l d be o b s e r v e d i n some  19.  RESULTS  Preliminary  experiments  P r e l i m i n a r y experiments u s i n g  goldfish  irradiated  with the cold quartz-mercury Mineralight  lamp were  qualitative  could  duced  and  indicated that  by u l t r a v i o l e t Radiations  blistering tips  an e f f e c t  g r e a t l y over t h a t  from | t o 2 h o u r s c a u s e d a  lasting  o f the h e a d  Susceptibility o f the c o n t r o l  and the  to' f u n g u s  fish  increased  k e p t i n the same  aquarium.  In s e v e r a l cases I n f e c t i o n o f the  ment a l s  heavy  was  irradiated  fish  enough t o  cause d e a t h .  i n the  o f s e v e r a l weeks.  The  b l i s t e r e d a r e a s on t h e h e a d spread g r a d u a l l y v e n t r a l l y . one  f i s h had  complete  Some o f t h e  changed  skin after  and t i p s  o f the  side of the  a r c b e l o n g i n g t o t h e Department  B l i s t e r i n g and  blistered which  fins  and weeks  from the g o l d c o l o r t o an almost  b l a c k on the d o r s a l  shortly after  latent  A t t h e end o f f o u r  p r o d u c e d s i m i l a r b u t more d r a s t i c fish.  a  t o the  b l a c k e n i n g b e g a n i n the  lateral  S e v e r a l experiments w i t h a strong mercury  experi-  showed i n c r e a s e d b l a c k e n i n g due  f o r m a t i o n of melanophores period  pro-  radiation.  of the d o r s a l r e g i o n s  o f the f i n s .  be  only  sluffing  the i r r a d i a t i o n .  on the  gold-  s k i n wer?e;.pr--o.duce:d  Fungus a p p e a r e d  a r e a s and. d e a t h o c c u r r e d  r e c o v e r e d from the fungus  quartz-  of Physics  effects  of the  line.  i n most c a s e s .  showed b l a c k e n i n g  on t h e Fish around  20.  the  b l i s t e r e d areas  One  fish  on  the  the  survived.  small  This  blood  clots  i n the  fins.  f i s h d e v e l o p e d dense pigment  d o r s a l head r e g i o n s ,  e y e s and  later  and  on  the  of the  operculum,  on the  tips  fins.  spread  t o the  v e n t r a l regions  One  o f the  most s t r i k i n g  The  behind  pigment  below the  lateral  lines.  i r r a d i a t i o n s was experimental  the  fish  decrease  sank t o the  where t h e y  remained with  were t h e s e  irradiated fish  days a f t e r  the  subjected  to  m o t i o n s and  fins  results  in activity  of  shown.  b o t t o m of the "folded".  these The  aquarium  I n no  case  o b s e r v e d to f e e d u n t i l  experiment took p l a c e .  similar handling  Control  showed a c t i v e  f e d immediately a f t e r  the  end  four  fish  swimming  of  the  experi-  ment. T h r e e e x p e r i m e n t s were c a r r i e d f i n g e r l i n g coho salmon a v e r a g i n g The 18  cm.  The  f i s h were p l a c e d  i n cross  outside,  air  j e t s were k e p t  i n length.  36  cm.  running  k e e p i n g the  was around  constant.  j a r d u r i n g the  x  cm.,  water c i r c u l a t i n g  temperature  i n the  (28  i n height).  w a t e r t o a d e p t h o f 15  i n a sink with  the  using  in a battery jar  s e c t i o n a l a r e a and  battery j a r , with  placed  5 cm.  out  Two  time the  fish  remained In i t . C o n t r o l s were r u n a glass plate placed jar.  i n a s i m i l a r manner, b u t  between the  I n e a c h c a s e the  RS  lamp and  sunlamp was  40  the cm.  with  battery above  the  21.  bottom o f the b a t t e r y j a r . In  one e x p e r i m e n t  29 out o f 30 f i s h , 12 h o u r s k i l l e d  irradiation  i n another experiment  a l l of the f i s h .  f o r 18 h o u r s  killed  i r r a d i a t i o n of  Controls  showed no  mortalities. Pish irradiated showed  so"/,fatriMtya f t e r  controls  i n the s i n k w i t h r u n n i n g w a t e r  1 week o f i r r a d i a t i o n , w h i l e  showed 2 d e a d out o f 3 0 . Power s o u r c e s were v a r i a b l e  but  the  results  do show t h a t  during  this  ultraviolet radiation  period,  h a s some  effect.  Chum s a l m o n eggs Eight  e x p e r i m e n t s were c a r r i e d  eggs o f the chum s a l m o n , b e g i n n i n g l i z a t i o n and a t t h e i n t e r v a l s until of  control  ation  are indicated  appeared by  and i r r a d i a t e d  that  the g l a s s p l a t e  produce a l e t h a l and  8.14x).  transmitted affect  III.  amount  Percentage  At l e s s e r  mortalities  after  irradi-  Prom t h e t a b l e i t  of u l t r a v i o l e t  o v e r t h e c o n t r o l s was effect  ferti-  i n Table I I I  eggs a t i n t e r v a l s  i n ^able  the small  6 hours a f t e r  indicated  t h e s u p p l y o f eggs r a n o u t .  out u s i n g the  sufficient to  at the higher doses doses the e f f e c t  transmitted  (37x, 18.5x,  o f the u l t r a v i o l e t  by t h e g l a s s p l a t e was n o t s u f f i c i e n t t o  the c o n t r o l s  as s t r o n g l y as the I r r a d i a t e d  fish.  Table I I I Date  Dosage  Total Sample Time i n Days  Nov. 28  37x  Dec. 1  18.5x  Dec. 4  8.14x  Dec. 6  5.14x  Dec. 6  3.4x  Dec. 7  1.7x  Dec. 9  1.29x  Dec. 17  Percentage mortalities of chum salmon eggs irradiated 6 hours after f e r t i l i z a t i o n and at various periods thereafter. Death expressed i n percentage of total sample 1  cont.  160  irrad.  188  cont.  152  irrad.  159  dont.  218  irrad.  217  Cont.  125  irrad.  125  100  100  10  irrad.  100  100  cont.  125  irrad.  2  4  5  6  8  9  10  11  12  14  Time i n Days 15  20  22  24  86.9 100 90.1 100 9.6 100 6.9  25.5  12.5  16.5  15  20  14.4  18.4  19.5  25.1  35.5  45.1  56.3  125  32  58.4  68  73.6  80.8  86.4  97.6  cont.  129  .77  irrad.  95  12.6  cont.  80  18.7  53.7  irrad.  80  98.7  100  cont.  23  2.27 63.1  17.1  24.3  31.05  25  22  21  42.25  36  7.7  13.1  23.2 25.5  80  87.3  97.9 56.3  99 58.5  98.7  0, - c o n t r o l Pig.  2  eggs  I - i r r a d i a t e d eggs  Graphs o f the p e r c e n t a g e m o r t a l i t i e s o f I r r a d i a t e d a n d c o n t r o l chum s a l m o n eggs over a p e r i o d o f days a f t e r dosages o f 3.14x, 2x, 1.7x, and 1.29x.  24  P i g . 2 shows g r a p h i c a l l y the p e r c e n t a g e m o r t a l i t y of the  the l o w e r i r r a d i a t i o n s irradiated Since  this period whether  eggs  f o r each  survived  None o f  to hatching.  the eggs were s e n s i t i v e t o h a n d l i n g a t  of development  the v i s i b l e  light  several handling  and s i n c e  over the c o n t r o l s  trans-  a f f e c t e d the  e x p e r i m e n t s were c a r r i e d  eggs were t r e a t e d  similarly  eggs,  a l l l i g h t was  except that  i t was n o t known  and n e a r u l t r a v i o l e t  m i t t e d by the g l a s s p l a t e ©ggs*  experiment.  t o I r r a d i a t e d and excluded.  r a n g e d f r o m 8 t o 14 p e r c e n t o v e r a p e r i o d Thus i t was c o n c l u d e d t h a t  the v i s i b l e  o u t . The  control  Mortalities of twenty  and near  days.  ultra-  violet  r a d i a t i o n s t r a n s m i t t e d t o t h e c o n t r o l s h a d some  effect  on the eggs  Sockeye  salmon  d u r i n g the e a r l y s t a g e s of  eggs  Six- s e t s u s i n g sockeye salmon  of i r r a d i a t i o n s eggs.  These  two d a y s b e f o r e t h e eggs became Intervals  development.  were c a r r i e d out experiments began  e y e d and t h e r e a f t e r a t  as shown i n T a b l e I V . Table  IV shows t h e d o s a g e ,  percentage m o r t a l i t y  of I r r a d i a t e d  Prom the 2x a n d 3x i r r a d i a t i o n s  total  sample  and c o n t r o l  eggs.  i t appeared t h a t the  c o n t r o l s were not a f f e c t e d w h i l e the i r r a d i a t e d suffered harmful effects  (Pig. 3).  t h e s e e x p e r i m e n t s were a c c i d e n t a l l y completion.  L a r g e r dosages  and  (35.5x,  eggs  Unfortunately interrupted before 16.2x, l l x ,  and  25.  3.43x) showed more s t r i k i n g f i s h b e i n g more s t r o n g l y  d i f f e r e n c e s , the  Irradiated  affected.  An i n t e r e s t i n g f e a t u r e o f t h e i r r a d i a t i o n o f December 20, 1948 was a lethal The  effect  the f a c t  thait a s w e l l  as p r o d u c i n g  o n the e g g s , h a t c h i n g was a c c e l e r a t e d .  u n t r e a t e d eggs i n t h e e x p e r i m e n t a l h a t c h e r y  commenced fish  to hatch  January  began to h a t c h  the i r r a d i a t i o n .  18, 1949.  on December  The  irradiated  2 1 , 1948, one d a y  A t t h e end o f the f i r s t  distortions  o f the i r r a d i a t e d  of body form  were a l l o w e d t o d e v e l o p 5 are photographs  fish  (Pig. 4 ) .  mortalities.  a number  fish  F i g u r e s 4 and  and c o n t r o l  s a l m o n a l e v i n s t a k e n 46 d a y s a f t e r  showed  These malformed  f o r some t i m e .  of i r r a d i a t e d  after  week h a l f o f  t h e eggs h a d begun t o h a t c h , b u t w i t h g r e a t Of t h e s u r v i v o r s  troughs  sockeye  irradiation  o f the  eggs. The development  control fish  i n pigmentation  y o l k , development fish  ( F i g . 5) showed more  of f i n s ,  and s i l v e r i n g , and g r o w t h .  advanced  a b s o r p t i o n of  The  irradiated  ( F i g . 4) showed d i s t o r t e d b o d y c o n t o u r s w i t h t h e  head depressed heart.  o v e r t h e y o l k s a c a t the l e v e l  The y o l k s a c was  absorbed  less  of the  r a p i d l y i n the  I r r a d i a t e d animals  and t h e f o r m a t i o n o f p i g m e n t  d e l a y e d 2 weeks.  Until  the time  i r r a d i a t i o n , pigmentation h e a d was could  lacking.  on t h e d o r s a l  The i r r a d i a t e d  swim o n l y i n c i r c l e s  of d e a t h  fish  was  75 days  after  r e g i o n s o f the were a c t i v e , b u t  s i n c e t h e i r b o d i e s were  distorted.  Table IV  Percentage mortalities f o r a period of days after irradiation  Total Date  Nov. 12  Nov. 13  Nov. 29 Dec. 2  Dec. 20  Deo. 30  Dosage  2x  3x  35.5x 16.2x  llx  3.43x  Experiments carried out using sockeye salmon eggs two days before eyeing and at intervals thereafter.  Sample Control  100  1 0  2 0  3 0  4 0  irrad  100  > 0  0  0  2  '  0  0  0  0  0  .45  2.6  control  40  irrad  40 220  0.  irrad  220  22.7  control  220  irrad  223  control  200  irrad  200  control  200  2  irrad  200  2  0 59.6  0  2.5 17.5  control  73.2  5 0  2  6.8  6 0  7 0  8 0  9  10  11  12  0  0  0  30  35  45  10  11  12  14  15  18  20  23  26  27  32  98.0  98.5  Experiments interrupted  8.6  7.7  11.3  100  91.8 0  0  .45  .90  3.15  1.8  86.5 86.9 89.7  98.6  96.4  3  4  48.5 59.5 8.5  75  4.5  6  87.5  89  31.0 29  100  34.0' 30  44.5  34.5 97.5n  100  Pig.  3 -  Graphs o f t h e p e r c e n t a g e m o r t a l i t i e s o f i r r a d i a t e d and c o n t r o l sockeye salmon eggs o v e r a p e r i o d o f d a y s a f t e r d o s a g e s o f 2x, 3x, 3.43x, l l x , 16.2x, and 35.5x.  Fig.  4 -  S o c k e y e s a l m o n a l e v i n s 46 days a f t e r i r r a d i a t i o n o f 35.5x. Mag. 5.5x.  an  30.  A similar 30,  1948, u s i n g  stimulated. vertebral  e x p e r i m e n t was  a d o s a g e o f 3.43x.  Of the e x p e r i m e n t a l  columns a n d c i r c u l a r  m e n t a t i o n was d e l a y e d slower  than  controls  carried  Hatching  fish,  ( P i g . 3 ) may  was  again  two showed  curved  swimming movements.  and a b s o r p t i o n  the c o n t r o l s .  out o n December  Pig-  o f the y o l k s a c was  Increased m o r t a l i t i e s  i n the  be a c c o u n t e d f o r by a n a t t a c k o f  fungus on t h e eggs. The  greatest m o r t a l i t y occurred  ments d u r i n g t h e h a t c h i n g p r o c e s s . first,  body f i r s t ,  normally fish  i n these  The f i s h  o r y o l k sac f i r s t ,  emerged h e a d  a s opposed t o the  h a t c h i n g f i s h w h i c h emerged t a i l  o n l y o c c a s i o n a l l y showed s i m i l a r  experi-  first.  Control  abnormalities i n  hatching. Sockeye salmon a l e v i n s E x p e r i m e n t s were c a r r i e d salmon a l e v i n s a v e r a g i n g began h a t c h i n g 27.  were u s e d . out  18, w i t h  These  fish  the p e a k on J a n u a r y  o f a l e v i n s a n d two c o n t r o l g r o u p s  4.  carried  The 0.5x a a d l x i r r a d i a t i o n s  out on February  11.  Since  a t u r e never r o s e above 3°C d u r i n g was  sockeye  The 2x, 4x, and 8x I r r a d i a t i o n s were  on F e b r u a r y  carried  20 cm. i n l e n g t h .  on January  F i v e groups  out u s i n g  the water  that time,  were  temper-  development  n o t v e r y ' . d i f f e r e n t a n d t h e two s e t s o f r e s u l t s  considered  together.  each s e t .  Table  C o n t r o l s were c a r r i e d  were  out f o r  V g i v e s t h e l i v e r e m o v a l s , R,  daily  Table V  Control ( l ) Total Sample Time  Q 1 2 3 4 5 6 7 8 9 10 11 12  13  14 15  16 17 18 19 20  a  22 23 24 25 26 27 28 29 30  5x  366 R  10 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2  0  c 0.28 0.29  0 0 1 0 1  0.89  0  0.91  0  0 1  0 0 0 0 0  0 2 0 0 0 2 0 0 -1 2 1 0 0 2 0 0 0 2 0 0 0 60 6  lx  341  M  0 1 0 0 0  Live removals, R, mortalities, M, and cumulative percentage mortalities, C, shown daily f o r a number of sockeye salmon alevins.  0.6  1.23 1.25 1.29 1.61 1.95 1.96 1.97  300 alive  R  303 C  R  0 0 0.61 2.13 10.73 11.73 13.97 18.13 31.76 43.67 50.32 58.07 65.81 75.0 81.04 83.22 85.09 86.33 86.79 87.83 89.12 89.72  5 2 2 2 2 2 2 2  M  5 0 2 0 2 2 2 5 2 28 2 3 2:: 7 2 13 2 43 2 35 2 22 2 23 2 232 27 2 17 2 5 2 4 2 2 2 0 2 1 2 2 2 0 0 0 2 4 0 2 2 0 0 1 2 0 0 0 2 0 0 0 55 269  91.72 92.41 93.01 93.40 94.06  17 alive  2 2 2 2 2 2 2  2 2 2 2 2 43  M  0 0 0 0 0 0 12 4.11 13 8.62 35 20.83 47 37.41 35 50.0 36 63.12 35 76.07 19 83.45 16 89.85 0 90.57 6 93.38 2 94.81 3 96.64 1 97.74 0 98.48 0 99.23 0 100.0 260  All -  c  Dead  .;R  5 2 2 2 2 2 2 2 0 19  2x  4x  8x  Control (2)  271  245  220  291  M  C  0 0 4 1.52 23 10.34 61 33.97 62 58.36 70 86.27 18 94.07 12 99.60 2 100.0 252  A l l Dead  R  5 2 2 3 2 2 2 18  M  C  0 0 7 2.94 30 15.67 106 61.37 42 80.08 42 99.12 0 100.0 227 A l l Dead  R  5 2 2 2 2 0 13  M  C  0 0 80 37.56 69 70.61 52 96.17 5 99.51 1 100.0 207  A l l Dead  R  M  10 0 2 2 2 1 2 2 2 0 2 0 2 0 2 5 2 4 24 14  C  0 0.71 1.08 1.82 1.83 1.84 1.86 3.74 5.24  253 Alive  32.  mortalities,  M,  and  c u m u l a t i v e p e r c e n t a g e d e a t h , C, f o r  each set o f i r r a d i a t i o n s . w i t h t h e 2x, was  f o r the  4x, and 0.5x  and  the  flow of water  heavy m o r t a l i t i e s  l x Irradiations.  -  of deaths f o r each sockeye a l e v i n s , i n each  end  The  t h r o u g h the c a r r y i n g  The  run (2)  experiments  trough caused  i n the trough.  calculated  sample  theoretical  o f c o n t r o l and  as compared t o t h e t o t a l  number  irradiated sample  used  experiment.  Experiment  T h e o r e t i c a l no. of deaths  C o n t r o l (1) 0.5x d o s e l . O x dose C o n t r o l (2) 2x d o s e 4x dose 8x dose The  attributable (Table  m e n t a l s were  T o t a l sample used  7.17 320.7 303 18.53 271 245 220  366 341 303 291 271 245 220  r e m o v a l s were compensated  would have been  made  that  o f 30 days when a d e c r e a s e  of a l l f i s h  Table VI  (1) was  8x i r r a d i a t i o n s , w h i l e C o n t r o l  were t e r m i n a t e d a t t h e in  Control  a slightly  The  not the removals  v a l u e s o b t a i n e d f o r the  significantly  there  g r e a t e r number o f d e a t h s  t o t r e a t m e n t had VI).  f o r since  different  been experi-  f r o m t h o s e of t h e  controls. In a l l the i r r a d i a t i o n s alevins,  the a c t i v i t y  o f the  carried  f i s h was  out w i t h t h e  definitely  affected,  33.  During the i r r a d i a t i o n s into the  the f a r t h e s t  the a l e v i n s  some o f the f i s h  weakened and sank t o the b o t t o m made o c c a s i o n a l  spasmodic  from t h e bottom  to the s u r f a c e  movements,  o c c a s i o n a l movements  These  remained  and active  Subsequent from  group  r e a c t i o n was  Some  less  When a b r i g h t  a similar  visibly  back  to d i e .  among t h e f o u r h o u r  In  swiftly  showed a g r a d a t i o n  among t h e c o n t r o l s .  shone on e a c h g r o u p  rising  only to f a l l  f i s h were o b s e r v e d t o be the f i r s t o b s e r v a t i o n s of a c t i v i t y  became  of the b a s k e t .  remain q u i e s c e n t f o r long p e r i o d s .  loss of activity  crowded  c o r n e r s o f the h o l d i n g b a s k e t s .  l o n g e r exposures  activity  actively  to great  light  was  noted.  This  i n the h i g h e r exposures  until  complete m o r t a l i t y o c c u r r e d / ' In less active mately  the two l o w e r i r r a d i a t i o n s , than the c o n t r o l s  15 days, when i t was  a f f e c t e d 'fish had d i e d place  experiment, exposure  for a period  observed that  and t h a t  among t h e r e m a i n d e r .  the f i s h  remained  of a p p r o x i -  the most  some r e c o v e r y h a d t a k e n  A t t h e c o n c l u s i o n o f the  the s e v e n t e e n r e m a i n i n g f i s h  showed the scene d e g r e e  o f t h e 0.5x  of a c t i v i t y  a s the  controls. Most  o f the f i s h  i n the h e a v i l y  g r o u p s were u n a b l e t o swim. and  spasmodic  jaw  i n respiration.  irradiated  The h e a r t b e a t became  slow  a n d many seemed u n a b l e t o move t h e l o w e r White  spots appeared  on the y o l k  Fig.  6a - Graph of d a i l y percentage oumclative mortal!ty  of sockeye salmon alevins irradiated with dosages of o.5x, lx» 2x, 4x, and 8x«  34. sac, and  the v i t e l l i n e v e i n s  appeared prominent  and d i s t e n d e d ,  i n many c a s e s a n a c c u m u l a t i o n o f s e r o u s  below the l a t e r a l  line  near the attachment  fluid  appeared  o f the y o l k  sac. Mortalities temperature  was  o c c u r r e d too r a p i d l y  t o o low t o o b t a i n  in  the g r o w t h r a t e .  was  r e t a r d e d about  Development  control  days  after  appeared  of the s i l v e r y  pigment  survived  6a) •  fish  Histological  studies  showed l i t t l e  irradiation.  effect  elastic  layer  tended.  The  cells  or b a s o p h i l i c  o f the d e r m i s goblet  on t h e f i r s t  cells  day  two  changes  o f the s k i n i n t h e  to the r a d i a n t  o f the s u p e r f i c i a l  two e o s i n o p h i l i c  of experimental  On the t h i r d  i n t h e e p i d e r m i s and d e r m i s  areas d i r e c t l y exposed nuclei  differences  studies  (A) S k i n and  significant  a week i n t h o s e f i s h w h i c h  t h e l o w e r e x p o s u r e s A^ig  Histological  and the water  energy.  rounded  The  up i n t o  one o r  g r a n u l e s and t h e f i b r o -  appeared  s w o l l e n and d i s -  were njore r o u n d e d  and t h e  p o r e s t o t h e e x t e r i o r were opened t o a w i d e r e x t e n t ( P i g . 6) t h a n t h o s e o f t h e c o n t r o l s On t h e f o u r t h d a y a f t e r a p p e a r e d more  numerous a n d c o u l d  b a s a l l a y e r s of the e p i d e r m i s were r e d u c e d i n number The  (Pig. 7 ) .  irradiation  be o b s e r v e d i n t h e  (Pig. 8).  The g o b l e t  or c o m p l e t e l y absent  s k i n o f t h e more h e a v i l y  the g r a n u l e s  irradiated  fish  cells  i n some a r e a s . (4x, 8 x ) was  35.  ragged ating  on i t s o u t e r s u r f a c e a s t h o u g h i t were (Pig. 9).  Lower d o s a g e s  (0.5x, I x , 2x) showed  g r a n u l a t i o n and r e d u c t i o n or l o s s did  degeneration  granules after  and  eosinbphilic  o c c u r r e d i n t h e h i g h e r .dosages  5 t o 8 days  (Fig. 11).  the d e r m i s became  Only  (Pig. 10).  of the s k i n w i t h the  a p p e a r a n c e o f numerous b a s o p h i l i c  of  of g o b l e t c e l l s , b u t  n o t show any s i g n s o f d e s q u a m a t i o n Severe  small fragments  w h i c h showed s e v e r e occurred before  desquam-  (4x, 8x, 2x)  The f i b r o e l a s t i c  layer  s w o l l e n on the 3 r d t o 4 t h d a y . o f i t c o u l d be  found  on t h o s e  fish  g r a n u l a t i o n of the e p i d e r m i s .  Death  s i g n s o f r e c o v e r y i n t h e 2x, 4x and 8x  dosages. Lower d o s a g e s  (0.5x, l x ) showed l e s s  g r a n u l a t i o n i n the e p i d e r m i s fibroelastic observed to  membrane.  set i n (Fig. 12).  a p p e a r a n c e o f r e c o v e r y was o f the e p i d e r m i s .  proliferation new  goblet  trols ules  the 1 3 t h The  first  t h e h e i g h t e n i n g of the b a s a l  cells  and t h e a p p e a r a n c e o f  cells. of s k i n from  the r a d i a t i o n s  appeared  t h e s i d e s o f t h e b o d y away  similar  (Fig. 7), with goblet c e l l s such  were  Subsequent r e c o v e r y i n v o l v e d  of e p i d e r m a l  Sections from  effects  These l a s t e d u n t i l  1 5 t h day when r e c o v e r y  cells  and l e s s d i s t e n s i o n o f t h e  The most s e v e r e  on the 8 t h d a y .  severe  as t h o s e f o u n d  t o t h a t o f the  present  i n the i r r a d i a t e d  and no areas.  con-  gran-  P i g . 6 - Skin of f i s h 3 days a f t e r an i r r a d i a t i o n of 4x. Mag. 440x.  P i g . 7 - Skin of c o n t r o l 7 day . began. Mag. 440x.  r the experiment  ok** F i g . 8 - Skin of f i s h 4 days a f t e r an of 2x. Ma^. 440x.  irradiation  Pig.  11 - S k i n o f f i s h 7 days a f t e r of 2x. Mag. 4 4 0 x .  an  irradiation  P i g . 12  - S k i n o f f i s h 13 days 0.5x. Mag. 440x.  a f t e r a dosage o f  40.  (B) O t h e r  tissues  -  T i s s u e s l y i n g deeper  f i s h d i d n o t show h i s t o l o g i c a l The  thyroid  and p i t u i t a r y  g l a n d s were s i m i l a r  c o n t r o l s b o t h i n development The  kidney  appeared  and mesonephric  not  constant.  and h i s t o l o g i c a l  unchanged i n both  regions.  were n o t p o s s i b l e  changes a f t e r  since  The digestive  t o the structure.  the p r o n e p h r i c cells  the t h i c k n e s s o f t h e s l i d e s  However, q u a l i t a t i v e  o r i n t h e number  irradiation.  Counts o f h e m o p o i e t i c  r e v e a l e d no c h a n g e s i n t h e n u c l e i cells  i n the  observations  of the hemopoietic  of r e d blood c e l l s  thymus g l a n d , g i l l s ,  t r a c t were n o t a f f e c t e d .  was  nervous  i n the kidney. system  and  41.  DISCUSSION  Ultraviolet  and  r a t e of  Hinricks  hatching  (1925) f o u n d  low  susceptibility  was  an a p p a r e n t t r a n s i t o r y  hatching. the  with less  trols.  T h e r e was  percentage  s t i m u l a t i o n with  hatching p e r i o d than  and  p e r i o d of  low  t i m e when t h e  at  s u s c e p t i b i l i t y was  e m b r y o n i c a x i s was  stages  hastening i s not  clear.  c h o r i o n of the  s i n c e the oped as  was  the  hatching.  t o be  that  produced  where l a r g e d o s e s  the  egg  to such  egg  not  an  c o n t r o l s which h a t c h e d  have  extent  disrupted this  apparently were not normally.  i n the h a t c h i n g process  that  membrane.  speeded as  a  effect  the r a d i a t i o n must  embryos w h i c h h a t c h e d  which occurred  found  mechanism o f  embryo m o v i n g w i t h i n t h e  Embryonic development  con-  development produced  The  I n some way  of  agree w i t h t h o s e  embryonic  of h a t c h i n g .  weakened t h e the  of  i n the  w e l l formed.  experiments r e p o r t e d here,  later  in  n o r m a l eggs i n e a c h l o t , s i n c e  These o b s e r v a t i o n s i n the  to  c o r r e l a t i o n between  o n l y t h e more n o r m a l embryos were c a p a b l e The  regard  exposed l o t s hatched  a significant  of hatched  of  intense r a d i a t i o n s there  More eggs i n t h e  e a r l y p a r t of the  that d u r i n g periods  up  fully The  devel-  mortalities  were h i g h e r  than  i n the c o n t r o l s . Mortalities The  i n eggs and  alevins  experiments c a r r i e d  out  show t h a t u l t r a v i o l e t  42.  has  a lethal  fish.  The  effect effect  since u l t r a v i o l e t l a y e r s of  The  believed  t o be  energy  i s primarily radiations  c a n t h e n be  ultraviolet  The  absorbed as a w h o l e  1935).  ( C l a r k , 1922).  ultraviolet,  e x p e r i m e n t s , was  a b s o r b e d by  especially  not  p r o t e c t e d by the  skin  the a c t i v a t i o n  and  o f the  of the e f f e c t s  of  i n the r e g i o n s used i n t h e s e  lying  skin,  Since  o v e r tiae y o l k s a c  i t was  probable that  t o an a p p r e c i a b l e e x t e n t .  emia p r o b a b l y p r o d u c e d t i s s u e s which  the  t h e h e a t i n g o f the t i s s u e s .  b l o o d . i n the v e s s e l s  heated  by  Thus one  v  the  b l o o d was  Giese,  s i n c e the b l o o d  g i v e n t o the o r g a n i s m  blood i s transformed to heat  the  thin  i n the corium i s  radiations.  P a r t of the energy  molecules  by  ( L a u r e n s , 1933;  r e g i o n absorbs the  visible  hatched  reaction  are absorbed  of g r e a t i m p o r t a n c e  of the  (Laurens,  a surface  l a y e r of c a p i l l a r i e s  through t h i s  and p a r t  s a l m o n e g g s and  s k i n and p r o t o p l a s m  1939).  flowing  on b o t h  This  c h a n g e s i n t h e b l o o d and  were h a r m f u l t o t h e  embryos and  was the hyperother  hatched  fish. However c e l l Blum  injury  (1945) b e l i e v e s t h a t  injury,  the c e l l s ,  the  responses.  and p h y s i o l o g i c a l  The  latent  radiations.  as a r e s u l t  e l a b o r a t e various substances which  physiological exposure  o c c u r s from  bring  period  r e s p o n s e s may  be  o f the about  between accounted  43.  for  by  assuming t h a t these  stances  are  e l a b o r a t e d , by  Increased has  physiologically active the  cell  concentration  sub-  a t a slow r a t e .  of a d i l a t o r  a c t u a l l y b e e n d e m o n s t r a t e d i n the  substance  s k i n and  blood  of  a n i m a l s t h a t have b e e n e x p o s e d t o u l t r a v i o l e t r a d i a t i o n . L e w i s and  Zotterman  (1926) s u g g e s t e d t h a t i t was  histamine-like  "H"  shown t h a t the  substance r e l e a s e d i s not histamine  the  term  substance.  H -substance  tt  has  M  Menkin  the  but  result  with  was  of c e l l  injury.  latter  capillary Edema was  d o s a g e s 2x,  of the was  y o l k sac  logical the  being  and  the  8x,  a direct  substance not as  is credited  result  alevins  especially  body w a l l .  and of  the  subocular  hatching  reaction  irradiated with  X-rays.  of 3 0 . d a y s and yolk sac,  lymph s i n u s .  treated f i s h  s e p a r a t i n g the  junction  A similar  body w a l l above t h e  i n the  irradiated  at the  (1948) who  days b e f o r e  e x a m i n a t i o n o f the valves  the  of l e u c o c y t e s  o b s e r v e d i n the  4x, and  observed a l o n g the  to  migration  edema showed a l a t e n t p e r i o d  and  i t s usefulness.  "Leucotaxine"  o b s e r v e d by. W e l a n d e r e t a l  l o w e r jaw  and  permeability.  c h i n n o k . s a l m o n a few The  has  " l e u c o t a x i n e " which i s e l a b o r a t e d  t i s s u e edema, the  with  evidence  c a p i l l a r y p e r m e a b i l i t y was  b r i n g i n g about the  Increased  l o s t much o f  (1938) p o s t u l a t e d t h a t  which caused i n c r e a s e d histamine  More r e c e n t  a  lymph and  was in  the  Histo-  showed damage venous  systems  44 I n s a l m o n embryos, r e s u l t i n g Thus the m o r t a l i t y c a n be p a r t i a l l y (1)  heat  accredited  i n e x t e n s i v e hemorrhage. o f the t o two  hyperemia caused  eggs and  young  salmon  factors:-  by  the a b s o r p t i o n of  the  rays. (2)  chemical  changes w h i c h  damaged c e l l s radiations  o c c u r i n the  as a r e s u l t  of the  blood  and  absorbed  causing c a p i l l a r y d i l a t a t i o n  and  edema.  Effects  of u l t r a v i o l e t  (A) iments light  skin  P r e l i m i n a r y experiments using goldfish  (1938) who  irradiated  skin  a t i o n produced  v e n t r a l l y and  lamp.  disappeared  of  Preliminary experultraviolet  T h i s was  i n producing  and p i g m e n t a t i o n  As was the a r e a s  on f i s h .  succeeded  w i t h a quartz-mercury  -  s e r v e d t o show t h a t  can have an e f f e c t  by H a r t  in  on f i s h  sluffing  of g o l d f i s h  The  Fukui  and  was  (1927) f o u n d  i n d u c e d b l a c k e n i n g , b e g i n n i n g on the  o f the  body and  ing that  to the areas below the regenerating tissues  g o l d f i s h had that  of the  fins,  lateral o f the  occurred  observed  agents  the t i p s  pigment-  weeks.  p o i n t e d out, pigmentation  laterally.  that  to produce  numerous  dorsal sides  with gradual  line.  first  to s p r e a d  He  also  b l a c k , g o l d or  these melanophores  spreadfound  white  t h e power o f p r o d u c i n g m e l a n o p h o r e s ,  the a b i l i t y  of  irradiated  n e c r o s i s and  i n two  greatest injury  suggested  and  appeared  45  t o be r e l a t e d t h e new  to the presence  7 times  humans p r o d u c e d cially  or t y r o s i n a s e i n  skin. Smith  X-rays  of t y r o s i n e  (1932) f o u n d  that  goldfish irradiated  t h e dosage r e q u i r e d new p i g m e n t  induced pigment took  to produce  i n 11 d a y s .  erythema i n  This  artifi-  55 d a y s t o d i s a p p e a r .  m e l a n o p h o r e s i n d u c e d by the X - r a y s were u s u a l l y d e l i c a t e and i r r e g u l a r than those found  with  The  more  on t h e f i s h  before  treatment. G o o d r i c h and T r i n k h a u s g o l d f i s h produced light.  (1929) f o u n d  p i g m e n t when i r r a d i a t e d w i t h  They c o n s i d e r e d t h a t the c a l i c o  invisibly with  that  colorless  d u r i n g development.  chromoblasts,  These m u l t i p l i e d  calico  ultraviolet  f i s h were s p o t t e d  singly or i n nests, and d i f f e r e n t i a t e d  i n d e p e n d e n t l y at i r r e g u l a r  i n t e r v a l s t o form  spots or  clusters  Ultraviolet  i n some  o f melanophores.  light  way  s t i m u l a t e d the p r o d u c t i o n o f i n c r e a s e d c l u s t e r s . M e l a n i n i s not a simple substance, but r a t h e r a mixture  o f pigments  b e e n shown t h a t  of I l l - d e f i n e d  composition.  the. s k i n c o n t a i n s a n enzyme  c a p a b l e o f c o n v e r t i n g t h e compound 3, 4 lalanine this  from  (dopase)  dihydroxypheny-  (dopa) i n t o m e l a n i n - l i k e m a t e r i a l ,  enzyme was a b s e n t  I t has  and t h a t  the s k i n o f a l b i n o s .  The  p r o b a b l e p r e c u r s o r o f 3, 4 d i h y d r o x y p h e n y l a l a n i n e i s tyrosine  (Hawk, O s e r a n d Summerson, E a r l i e r workers t r i e d  1947).  t o e x p l a i n the a c t i o n o f  46.  ultraviolet  on s k i n , p r o d u c i n g p i g m e n t , a s a  o x i d a t i o n in-which  t y r o s i n e was d i r e c t l y  3, 4 d i h y d r o x y p h e n y l a l a n i n e , i n the presence tyrosinase. of melanin  photo-  oxidized to  the o x i d a t i o n t a k i n g p l a c e  of u l t r a v i o l e t  even i n the absence o f  More r e c e n t l y t h e a c t i o n o f the p r o d u c t i o n i n the s k i n has been e x p l a i n e d i n a  d i f f e r e n t manner.  The 3, 4 d i h y d r o x y p h e n y l a l a n i n e i s  now b e l i e v e d t o be t r a n s p o r t e d t o t h e s k i n f r o m parts  slightly  other  o f the body, and i n t h e s k i n i s o x i d i z e d t o  by dopase.  This l a t t e r  between the I r r a d i a t i o n melanin  (Blum, The  f i s h used  melanin  theory e x p l a i n s the l a t e n t p e r i o d o f t h e s k i n and t h e a p p e a r a n c e o f  1945). appearance  i n these  o f new m e l a n o p h o r e s i n the g o l d -  experiments  was d e l a y e d f r o m  2 to 3  weeks and i n c r e a s e d i n amount f o r p e r i o d s o f f r o m  3 to 5  weeks. Fungus w h i c h a t t a c k e d the i r r a d i a t e d f i s h first  on t h e s l u f f i n g and damaged a r e a s  later  spread to t h e surrounding  a s a p r o p h y t i c organism tissues f i r s t  and l a t e r  causing t h e i r  death.  (B)  Alevin  areas.  o f the s k i n and T h i s fungus i s  w h i c h a t t a c k s d e a d or damaged  sockeye  spreads  to the surrounding  salmon experiments  (1928) d e s c r i b e d t h e h i s t o l o g i c a l skin:-  appeared  changes of  -  tissues,  Laurens irradiated  ~  The changes o c c u r b r i e f l y a s f o l l o w s : the vessels become d i l a t e d w i t h i n f i l t r a t i o n o f the e p i t h e l i a l  47.  l a y e r a n d an a b n o r m a l t h i c k e n i n g and d a r k e n i n g . F o l l o w i n g i n t e n s e r a d i a t i o n ther.e i s s e r o f i b r i n ous o r c e l l - r i c h , o f t e n h e m o r r h a g i c , exudation. I f t h e i r r a d i a t i o n i s s e v e r e enough t h e c o l l a g e n o u s t i s s u e becomes s w o l l e n and homogeneous, t h e e p i t h e l i a n s w o l l e n and l o o s e n e d , i n f i l t r a t e d and b l i s t e r e d , the l o s s o f c o n t i n u i t y o c c u r r i n g a t the b o u n d a r y b e t w e e n t h e g r a n u l a r and h o r n y l a y e r s . If irradi a t i o n i s q u i t e i n t e n s e t h r o m b i f o r m i n the v e s s e l s of the c u t i s . I r r a d i a t i o n of the e a r of r a b b i t s w i t h a p o w e r f u l C a r c lamp r e s u l t s i n d e g e n e r a t i o n and n e c r o s i s o f the e p i t h e l i u m , b l i s t e r f o r m a t i o n , s t a s i s , f i b r i n o u s e x u d a t i o n t o t h e s u r f a c e and t i s s u e spaces. In a d d i t i o n there i s a r e l a t i v e l y s l i g h t w h i t e c e l l i n f i l t r a t i o n , m a i n l y o f lymphoc y t e s , with plasma c e l l f o r m a t i o n . Finally a marked p r o d u c t i o n of f i b r o b l a s t s and t h e f o r m a t i o n o f new v e s s e l s and r e g e n e r a t i v e p r o c e s s e s i n t h e epithelium.  The  results  somewhat t h e mammals.  in  skin.  i n the e p i d e r m a l  Since  fish  do  and  fish  skin  lower  layers  o f the  and clearly  fibroelastic  layers  n o t have a h o r n y l a y e r  i n the  the r a d i a t i o n s were p r o b a b l y a b l e t o epidermis,  penetrate  causing severe  damage  that region. The  damage t o the g o b l e t c e l l s  3 days a f t e r i r r a d i a t i o n p r o b a b l y had osmotic  regulation  surface  of the  osmotic  regulation,  damaged. chlorirle, off  resemble  changes i n the s k i n o f human b e i n g s  changes  epidermis, t o the  experiment  S t u d i e s o f s e c t i o n s of t h e  indicated of the  obtained i n this  of t h e f i s h .  Loss  some e f f e c t  especially  i f the  t o the  on  the  o f mucus on t h e  s k i n i s b e l i e v e d t o h a v e an  F i s h i n f r e s h water lose  or i f the  which occurred 2 to  effect  on  outer  the  underlying tissue  salts,  is  especially  s u r r o u n d i n g medium when t h e mucus i s w i p e d  s k i n i s damaged  (Krogh,  1939).  This loss  of  48.  osmotic death  c o n t r o l was  of t h e The  either  probably  a contributing factor  changes which o c c u r r e d i n the  or by the n u c l e u s .  s k i n may  r a d i a n t energy Giese  that  were more s t r o n g l y I n h i b i t e d i n growth and w a v e l e n g t h 2804 A, protoplasm.  A w a v e l e n g t h o f 2654 A,  growth i n i t i a l l y ,  In  bacteria  the  was  immediate  Paramecia by  the  found  Prom t h i s  of u l t r a v i o l e t  nucleus.  I f the  less  r e c o v e r y p e r i o d was  growth.  inhibition prolonged.  t o be he  was  more  concluded more on  Recovery of the  n u c l e u s was  the  the  cytoplasm  harmed r e c o v e r y  was  rapid.  the  nucleus  the  irradiated  irradiation. and  dosages  The  secondly  d o s a g e s showed no ( l x , 0.5x)  the b a s a l c e l l s . p r o t e c t i o n by not  the  effect  t h a n on t h e  In by  by  the r e g i o n o f maxi-  t h e w a v e l e n g t h 2654 A was  more r a p i d .  less  but  in inhibiting  cytoplasm  due  the r e g i o n of maximal a b s o r p t i o n o f  of  that  be the  killed  a b s o r p t i o n of the n u c l e o p r o t e i n s , caused  effective  by  (1939) f o u n d  u s i n g m o n o c h r o m a t i c beams o f u l t r a v i o l e t  mal  the  fish.  t o the a b s o r p t i o n of the  protoplasm  to  the  s k i n the whole c e l l g r a n u l a t i o n appeared  i n the  cytoplasm.  r e g e n e r a t i o n o f the  The  affected  first  i n the  higher  damaged s k i n .  Lower  showed r e g e n e r a t i o n o c c u r r i n g f i r s t These c e l l s  were p r o b a b l y  overlying cells  damaged b e y o n d r e c o v e r y .  experiments  was  were i n the  The  so t h a t t h e  afforded n u c l e i were  wavelengths used  r e g i o n which  Giese  in  (1939)  In  these  found  49.  most a b s o r b e d by The ized,  since  radiation  the  observed e f f e c t areas  of  showed no  produced i n these e a r l i e r , produce to the  rest  o r g a n i s m as Effect  of  after  ation  of  and  not  "injury"  f r o m the  changes.  areas,  as  was  was  by  the  localdirect  The  injuries  pointed  substances which are  tissues  the  b l o o d and  on  other  out  transported  affect  the  of  tissues  flocculent  c o a g u l a t i o n of Similar  were a t t r i b u t e d fact  on  c o n s i s t e n t w i t h the measured the  Their findings  the  same  tissues  yolk  the  surface  denatur-  i n .the  yolk  Welander et  al  X-rays,  cause.  t h a t the  layers  observations  of  a layer  of  i n the  ultraviolet tissue.  Bachem and  of  b o d y were  skin  to 1.0  X-rays  rays  This Reed  was  (1931)  ultraviolet. mm.  thick  u l t r a v i o l e t w h i l e v i s i b l e and  r a d i a t i o n s were more p e n e t r a t i n g . h i g h energy are  by  t r a n s p a r e n c y o f human s k i n  the  to  contained  l y i n g deeper  indicates  showed t h a t  a b s o r b e d most o f  proteins  the  chinook salmon a l e v i n s w i t h  to  that  v i s i b l y affected  the  s p o t s were o b s e r v e d  treating  infrared  because of  their  more p e n e t r a t i n g t h a n u l t r a v i o l e t r a y s  t h e i r primary effect d e e p e r i n the  spots  i r r a d i a t i o n were p r o b a b l y due  were a b s o r b e d i n the  who  ultraviolet  skin protected  localized  the  the  histological  appearance  (1948) a f t e r  The  of  a whole.  material.  and  the  ultraviolet  The sac  protoplasm.  body  is  on  the  (Bonham e t  organs a l , 1948;  and  tissues  lying  Welander et  al,  and  50. 1948).  The s u p e r f i c i a l  layers  were n o t v i s i b l y harmed, s i n c e appreciably  absorbed  i n those  such as muscle the r a d i a t i o n s  layers.  and  skin  were n o t  51,  SUMMARY 1. to  Ultraviolet  3100  of wavelengths  f r o m 2800 A  A causes l e t h a l or h a r m f u l e f f e c t s  sockeye lings  light  salmon  and 2.  on chum  eggs, a l e v i n s o c k e y e s a l m o n ,  coho  salmon  eggs  i n l a t e r stages of d e v e l o p -  ment were i n d u c e d t o h a t c h a month e a r l i e r t h a n developing f i s h .  Slowing of development  a t i o n were o b s e r v e d and  fish  finger-  goldfish. Sockeye  3.  and  abnormal  P i g m e n t a t i o n and  s k i n and  increased  and  normal  pigment-  body f o r m s were  produced.  s l u f f i n g were p r o d u c e d i n g o l d -  s u s c e p t i b i l i t y t o fungus  was  observed. 4. violet  Histological radiation  primarily skin.  sockeye salmon  e f f e c t was  localized.  i n t h e body showed no  5. but  on a l e v i n  The  actual  of u l t r a -  revealed  (a)  heat  (b)  development  and  of the  changes.  may  not  have been  determined, responsible:-  of t o x i c  s u b s t a n c e s i n the  tissues.  of osmotic r e g u l a t i o n  injuries.  changes  tissues  hyperemia.  Injured loss  Organs  c a u s e o f d e a t h was  a c o m b i n a t i o n of f a c t o r s  (c)  of the e f f e c t s  i n t h e e p i d e r m i s and f i b r o e l a s t i c - l a y e r s  The  deeper  studies  due  to  skin  52.  ACKNOWLEDGMENTS  The a u t h o r w i s h e s t o e x p r e s s h e r t h a n k s t o D r . W. A. Clemens a n d t h e members of t h e s t a f f o f the Department o f Z o o l o g y , 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 , where t h i s work was u n d e r t a k e n . S p e c i a l t h a n k s a r e due t o D r . W. S. H o a r , u n d e r whose g u i d a n c e t h i s work was c a r r i e d o u t and w i t h o u t whose h e l p f u l c r i t i c i s m s much w o u l d have been l e f t undone. Mr. J . Mason, Mr. I . B a r r e t t , Mr. R. W. R a d c l i f f e , Mr. P. P a u l a n d Mr. H. B e l l were most h e l p f u l i n p r o v i d i n g eggs a n d f i s h , b u i l d i n g equipment n e c e s s a r y , and p r e p a r i n g i l l u s t r a t i o n s . Many t h a n k s a r e d u e t o them f o r t h e . i r a s s i s t a n c e . V e r y s p e c i a l t h a n k s a r e due t o my p a r e n t s , whose g e n e r o s i t y and i n t e r e s t have e v e r b e e n an I n s p i r a t i o n f o r my work.  53.  LITERATURE CITED  Bachem A. and C. I . Reed, 1948. t h r o u g h human s k i n . Am.  The p e n e t r a t i o n o f l i g h t J o u r n . P h y s . , 97; 8 6 - 9 1 .  Blum, H.P., 1945. The p h y s i o l o g i c a l e f f e c t s on man. P h y s . Rev., 25: 483-530.  of  sunlight  Bonham, K., L . R. D o n a l d s o n , R. P. P o s t e r , A. D. W e l a n d e r and A. H. Seymour, 1948. The e f f e c t o f X - r a y on m o r t a l i t y , w e i g h t , l e n g t h , and c o u n t s o f e r y t h r o c y t e s and h e m a t o p o i e t i c c e l l s i n t h e f i n g e r l i n g C h i n o o k s a l m o n , O n y c o r h y n c h u s t s c h a w y t s c h a Walbaum. Growth, 2: 107-121." B u r g e , W.E., 1917. The a c t i o n o f u l t r a v i o l e t r a d i a t i o n i n k i l l i n g l i v i n g . c e l l s such as b a c t e r i a . Am. J o u r n . P h y s . , 43: 429-432. B u r g e , W. E . , W. R. F i s c h e r a n d A. G. N e i l , 1916. Dest r u c t i o n o f hormones, p r o e n z y m e s , and enzymes b y ultraviolet radiation. Am. J o u r n . P h y s . , 40: 426-436.. I n H i n r i c k s , M. A., 1925. M o d i f i c a t i o n o f d e v e l o p m e n t on the b a s i s of d i f f e r e n t i a l s u s c e p t i b i l i t y t o r a d i a t i o n . I Fundulus h e t e r o l i t i c u s and u l t r a v i o l e t . r a d i a t i o n . J o u r n . Morph., 4 1 : 239-268. Clark,  J . H., 1922. The p h y s i o l o g i c a l P h y s . Rev., 2: 277-307.  action  of  light.  F u k u i , K e n ' I c h i , 1927. On t h e c o l o r p a t t e r n p r o d u c e d b y v a r i o u s agents i n the g o l d f i s h . F o l i a . Anat. Japon., 5: 257-302. G i e s e , A. C , 1939. U l t r a v i o l e t r a d i a t i o n and c e l l d i v i s i o n . 1. E f f e c t s o f X 2654 and >v 2804 u p o n Paramecium caudatum. J o u r n . C e l l . Comp. P h y s . , 13: 139-150. G o o d r i c h , H. B. and J . P. T r i n k h a u s , 1939. The d i f f e r e n t i a l e f f e c t o f r a d i a t i o n s on t h e M e n d e l i a n p h e n o t y p e s o f the g o l d f i s h , C a r a s s i u s a u r a t u s . B i o l . B u l l . , 77: 188-195. Harris,  Hart,  F. I . and H. S. H o y t , 1917. The p o s s i b l e o r i g i n of the t o x i c i t y o f u l t r a - v i o l e t l i g h t . Science, 46: 318-320.  J . L . , 1938. ' Can s u n b u r n a f f e c t f i s h i n h a t c h e r y ponds? F i s h . R e s . Bd. Can. P r o g . R e p t . P a c , ( 3 5 ) : 6.  54.  Hawk, P. B», B. L. Oser and Wm. H. Summerson, P r a c t i c a l p h y s i o l o g i c a l chemistry, x i v + 1 2 t h ed., B l a k i s t o n Co., Philadelphia.  1947. 1-1322,  H i n r i c k s , M. A., 1925. M o d i f i c a t i o n of d e v e l o p m e n t on the b a s i s o f d i f f e r e n t i a l s u s c e p t i b i l i t y t o radiation. I F u n d u l u s h e t e r o l i t i c u s and u l t r a v i o l e t radiation. J o u r n . Morph., 41: 239-268. K r o g h , A., 1939. Osmotic r e g u l a t i o n 1-242, C a m b r i d g e .  In  aquatic  animals,  L a n d o l t , H. and R. B o r n s t e i n , 1923. c h e m i s c h e t a b e l e n , 2: 896.  Physikalisch-  L a u r e n s , H., 1928. The p h y s i o l o g i c a l P h y s . Rev., 8: 1-91.  effects  of  L a u r e n s , H., 1933. The p h y s i o l o g i c a l e f f e c t s o f e n e r g y , 1-610, C h e m i c a l C a t a l o g u e Co., New L a u r e n s , H., 1935. Photochemistry i n medicine: a outline. Symp. Q u a n t . B i o l . , 3: 277-295.  radiation.  radiant York. general  L e w i s , T. and Y. Z o t t e r m a n , 1926. H e a r t , 13-: 203. I n Blum, H. P., 1945. The p h y s i o l o g i c a l e f f e c t s o f on man. P h y s . Rev., 25: 483-530. L u c k i e s h , M., 1946. e r y t h e m a l and No s t r a n d Co..,  The a p p l i c a t i o n o f germicidal, i n f r a r e d e n e r g y , v i i i + 1-463, D. New York.  sunlight  Van  M a y e r s o n , H. S., 1935. The e f f e c t s o f r a d i a t i o n on c i r c u l a t i o n , b l o o d and m e t a b o l i s m w i t h p a r t i c u l a r r e f e r e n c e t o - g r o w t h , b a s a l m e t a b o l i s m and t h e thyroid gland. Symp. Q u a n t . B i o l . , 3: 299-311. M e n k i n , V., 1938. P h y s . Rev.,  The r o l e of i n f l a m m a t i o n i n 18: 366-418.  immunity.  P o h l e , E. A., 1926. S t a n d a r d i z a t i o n of u l t r a - v i o l e t r a y s . The u l t r a - v i o l e t u n i t . J o u r n . Am. Med. A s s o c . , 86: '818-820. Smith,  G. M., 1932. M e l a n o p h o r e s i n d u c e d by X - r a y as compared w i t h t h o s e e x i s t i n g i n the p a t t e r n s as s e e n I n C a r a s s i u s a u r a t u s . B i o l . B u l l . , 63: 484-491.  W e l a n d e r , A. D., L. R. D o n a l d s o n , R. P. P o s t e r , K. Bonham and"A. H. Seymour, 1948. The e f f e c t s o f R o e n t g e n r a y s on the embryos and l a r v a e o f t h e Chinook s a l m o n . Growth, 12: 203-242.  55.  Weitz,  C. E . , 1946. General E l e c t r i c B u l l . LD1, T o r o n t o .  lamp  bulletin.  ABSTRACT  Ultraviolet 3100 A,  light  o f w a v e l e n g t h s f r o m 2800 A t o  g e n e r a t e d by a G e n e r a l E l e c t r i c  used to i r r a d i a t e  eggs  and  young  RS  fish.  sunlamp,  were  The d o s a g e s  were  s t a n d a r d i z e d b y means o f a p h o t o c h e m i c a l r e a c t i o n i n w h i c h i o d i n e was  liberated  of intense  light.  with the s t a r c h  from h y d r i o d i c This free  a c i d u n d e r the  iodine  gave a b l u e  was  auratus) produced d e f i n i t e  observed to b l i s t e r  susceptibility controls,  and  and s l u f f  The  o f t h e chum s a l m o n  of the  i n t h e number o f m e l a n o p h o r e s fish after  a  latent  c u r v e s were d e t e r m i n e d f o r the ( O n y c o r h y n c h u s k e t a ) and the  (Onycorhynchus  stages o f development  sockeye  month a h e a d  o f t h e c o n t r o l s were a b n o r m a l  The  so r a p i d l y  effect  a l e v i n s which hatched a  The v e r t e b r a l c o l u m n was  the r e g i o n o f the h e a r t ,  i n the  produced a s t i m u l a t i n g  of h a t c h i n g .  not u t i l i z e d  eggs  nerka).  on t h e r a t e  pects.  regions,  over that  E x p e r i m e n t s u s i n g s o c k e y e s a l m o n eggs later  skin  o f 2 t o 3 weeks. Mortality  salmon  effects.  t o f u n g u s became I n c r e a s e d an i n c r e a s e  goldfish  i n the exposed  o c c u r r e d i n some o f t h e i r r a d i a t e d period  coloration  test.  P r e l i m i n a r y experiments using the (Carassius  Influence  i n several  res-  c u r v e d downward o v e r  g r o w t h was  retarded,  as t h e c o n t r o l s  the yolk  was  and p i g m e n t a t i o n  was  delayed.  More m o r t a l i t i e s  the i r r a d i a t e d  occurred during hatching In  t h a n i n the c o n t r o l  Sockeye s a l m o n a l e v i n s were i r r a d i a t e d occurred  fish.  a v e r a g i n g 20  shortly after hatching.  in irradiations  l a s t i n g from  cm.  Heavy  i n length  losses  15 m i n u t e s t o 4  hours.  Histological  s t u d i e s r e v e a l e d changes i n the e p i d e r m i s  fibroelastic  layers  r e g i o n s which had elastic  sluffing  a latent  T i s s u e s u n d e r the  show v i s i b l e  changes.  and  l o s s of  goblet c e l l s . outer c e l l s  i n a s m a l l number i r r a d i a t e d w i t h t h e  dosages  subse-  Lower d o s a g e s  o f the  p e r i o d of 2 t o 3 days.  with  t o the fibro-  Larger  degeneration of epidermal c e l l s  showed d e g e n e r a t i o n o f t h e after  s k i n w h i c h were l o c a l i z e d  been e x p o s e d .  l a y e r d i d not  showed s e v e r e quent  o f the  and  epidermis  Recovery lowest  occurred  dosage  about I  12  t o 13  days a f t e r  irradiation.  Mortalities  c o u l d have b e e n c a u s e d  1.  heat  2.  c h e m i c a l c h a n g e s o c c u r r i n g i n the  result 3.  hyperemia caused  of c e l l  decrease  injuries  to  injury  by  caused  by:-  a b s o r p t i o n of the r a y s .  by  the  b l o o d as  radiations.  i n osmotic r e g u l a t i o n caused  the  skin.  by  a  

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