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Effects of low levels of 2, 4-dichlorophenoxyacetic acid on the uptake, translocation, and incorporation… Etter, Harold MacDonald 1966

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EFFECTS ON THE  OF  LOW L E V E L S  UPTAKE, BY  OF  2,4-DICHLOROPHENOXYACETIC  TRANSLOCATION,  THE  BEAN P L A N T  AND  INCORPORATION  (PHASEOLUS  OF  VULGARIS)  by  HAROLD B.S.A.,  University  M.Sc.,  A  THESIS THE  Oregon  of  IN  REQUIREMENTS DOCTOR  ETTER  British  State  SUBMITTED  in  M.  University,  PARTIAL FOR THE  OF  1961  Columbia, 1963  FULFILMENT DEGREE  OF  OF  PHILOSOPHY  the Department  of  BOTANY  We a c c e p t required  THE  this  thesis as conforming  to  standard  UNIVERSITY  OF May,  B R I T I S H COLUMBIA  1966  the  ACID P  3 2  iV  In p r e s e n t i n g t h i s t h e s i s  in p a r t i a l  f u l f i l m e n t of the requirements  f o r an advanced degree a t the U n i v e r s i t y of B r i t i s h Columbia, 1 agree t h a t the L i b r a r y s h a l l make i t f r e e l y avai1able .for r e f e r e n c e and study. I f u r t h e r agree t h a t p e r m i s s i o n f o r e x t e n s i v e c o p y i n g o f t h i s t h e s i s f o r s c h o l a r l y purposes may by h i s r e p r e s e n t a t i v e s .  be g r a n t e d by the Head o f my Department o r I t i s understood t h a t c o p y i n g o r p u b l i c a t i o n  of t h i s t h e s i s f o r f i n a n c i a l g a i n s h a l l not be a l l o w e d w i t h o u t my w r i t ten  permission.  Department o f  Botany  The U n i v e r s i t y o f B r i t i s h Vancouver 8 , Canada Date  May,  1966  Columbia  The U n i v e r s i t y of B r i t i s h Columbia FACULTY OF GRADUATE STUDIES PROGRAMME OF THE FINAL ORAL EXAMINATION FOR THE  DEGREE OF  DOCTOR OF PHILOSOPHY of HAROLD MACDONALD ETTER B.S.A., The U n i v e r s i t y of B r i t i s h Columbia 1961 3  M.Sc, Oregon State U n i v e r s i t y , 1963 TUESDAY, JUNE 7, 1966 AT 3:30 P.M. IN ROOM 3332, BIOLOGICAL SCIENCES BUILDING COMMITTEE IN CHARGE Chairman: I . McT. Cowan J . J . R. Campbell, D. P. Ormrod G. M. Tener Superviser:  G. H. N. Towers E. B. Tregunna D. J . Wort D. J . Wort  External Examiner: R. T. Wedding Professor, Biochemistry U n i v e r s i t y of C a l i f o r n i a Riverside, California  EFFECTS OF LOW. LEVELS OF 2,4-DICHLOROPHENOXYACETIC ACID ON.THE UPTAKE TRANSLOCAHON AND INCORPORATION OF P BY THE BEAN PLANT (PHASEOLUS VULGARIS) s  3 2  ABSTRACT A study was c a r r i e d out t o gain information, which could be used to describe the mechanism whereby 2 4-D a f f e c t s the growth of i n t a c t plants„ 5  Fourteen-day-old bean plants (Phaseolus v u l g a r i s ) growing i n a phosphate-free or complete n u t r i e n t s o l u t i o i were sprayed t o d r i p with 5 or 50 ppm 2,4-D. At various times up to 70 hours a f t e r spraying, the roots were immersed i n a complete n u t r i e n t s o l u t i o n containing P-^2. Following an incorporation period of 2-9 hours, the plants were harvested, and separated i n t o r o o t s stems plus petioless and leaves. Acid soluble a c t i v i t y , acid i n s o l u b l e a c t i v i t y , and the d i s t r i b u t i o n of a c t i v i t y w i t h i n the soluble f r a c t i o n as revealed by paper chromatography were determined f o r each organ. 3  In general the treatments had r e l a t i v e l y small e f f e c t s on phosphorus uptake, t r a n s l o c a t i o n , or incorpora t i o n i n t o organic compounds. I n i t i a l increases i n the rate of P-^2 uptake were not maintained and were followed by the same or l e s s uptake as c o n t r o l s . The rate of upward t r a n s l o c a t i o n was reduced and l e s s a f f e c t e d by the age of-the p l a n t s , as compared t o c o n t r o l s , a f t e r treatment with 5 ppm 2,4-D. The higher concentration did not a l t e r the d i s t r i b u t i o n of a c t i v i t y among the organs. While the lower l e v e l did not change the rate of incorporation i n t o the t o t a l acid i n s o l u b l e f r a c t i o n of the plant, the growth-inhibitory l e v e l reduced the a c t i v i t y i n t h i s f r a c t i o n by 14-22%.. At 5 ppm there was a trend toward more incorporation i n t o i n s o l u b l e compounds i n the roots and l e s s i n the leaves, but no cons i s t e n t change i n the stems. The composition of the soluble f r a c t i o n s showed no uniform v a r i a t i o n s from controls at e i t h e r concentration. The r e s u l t s i n d i c a t e d a pattern whereby the balance i n p32 d i s t r i b u t i o n between the leaves and roots was upset i n favor of the roots by f o l i a r a p p l i c a t i o n of 5 ppm 2,4-D, but not 50 ppm. Both stimulatory and i n h i b i tory l e v e l s of 2,4-D produced responses which appear to be r e l a t e d to the synthesis of acid i n s o l u b l e compounds i n the roots and leaves. The major p o r t i o n of the i n soluble a c t i v i t y i n these t i s s u e s was a l s o ethanol i n s o l u b l e and i s believed t o be RNA  Actigrapb scans of p i c r i c acid chromatograms i n d i cated more incorporation of a c t i v i t y i n t o organic compounds (nucleotides and sugar phosphates) i n the l e a f and stem t i s s u e s than i n the roots. Plants grown i n complete n u t r i e n t p r i o r to exposure to p32 took up, translocated, and retained l e s s phosphate than those grown i n phosphate-free nutrient,. Also, incorporation i n t o soluble organic compounds was suppressed. 3  The only r a d i o a c t i v e compound i n xylem exudates from plants 10 hours a f t e r t h e i r i n i t i a l exposure to p32 was orthophosphate. GRADUATE STUDIES F i e l d of Study: Plant Physiology Plant Physiology I , Anabolism, Growth D. J . Wort Plant Physiology I I , Catabolism, Water r e l a t i o n s D. J . Wort Plant Physiology Recent Advances D. .J. Wort Botany Graduate Seminar Faculty Other Studies: Outlines of Biochemistry Faculty Biochemistry of Proteins W. .J..Polglase Biochemistry of the Nucleic Acids G. M. Tener Biochemistry of Amino Acids S. H. Zbarsky Biochemistry of Carbohydrates W. J . Polglase Biophysics ---C. P. S. Taylor Laboratory Methods' A. Gronlund  i i  HAROLD M. ETTER. EFFECTS OF LOW LEVELS OF 2,4-DICHL0R0PHEN0XYACETIC ACID^QN THE UPTAKE, TRANSLOCATION, AND INCORPORATION OF BY THE BEAN PLANT {PHASEOLUS VULGARIS) Supervisor: D. J . Wort. "'< y  ABSTRACT*..-''  I A  study  describe  was c a r r i e d o u t t o gain  t h e mechanism whereby  Fourteen-day-old phosphate-free  bean  5 o r 50 p p m 2 , 4 - D .  information  2,4-D a f f e c t s  plants  o r complete  ,  At various  vulgaris)  solution  times  could  the growth  (Phaseolus  nutrient  which  were  of  intact  growing  sprayed  u p t o 70 h o u r s  be u s e d  plants.  in a  to drip  after  to  with  spraying,the  32 roots  were  Following and  immersed  an i n c o r p o r a t i o n  separated  activity, within  acid  determined  rus  into  roots,  stems  fraction  f o r each  general  uptake,  period  nutrient o f 2-9  plus  insoluble activity,  the soluble  In  in a complete  solution  hours,  petioles,  containing P  the plants and leaves.  and the d i s t r i b u t i o n  as revealed  by paper  were  harvested  Acid  of  soluble  activity  chromatography  were  organ.  the treatments  translocation,  had r e l a t i v e l y  or incorporation  small  into  effects  organic  on phospho-  compounds.  In-  32 itial  increases  followed  was reduced  to controls,  centration  the lower  total  acid  trend  level  insoluble  the activity  toward  more  uptake  uptake  and less  after  d i d not alter  While  reduced  of P  b y t h e same o r l e s s  translocation compared  in the rate  affected with  the distribution  d i d n o t change  in this  fraction into  of  and were upward  by t h e a g e o f t h e p l a n t s , a s  of activity  the rate  by  The r a t e  5 ppm 2 , 4 - D .  of the plant,  incorporation  not maintained  as controls.  treatment  fraction  were  of  The h i g h e r  among  the organs.  incorporation  into the  the growth-inhibitory  14-22%.  A t 5 ppm t h e r e  i n s o l u b l e compounds  con-  level was a  in the roots  and  less  in  the  leaves,  composition  of  controls  either  at  the  but  soluble  no  consistent  fractions  change  showed  no  in  the  uniform  stems.  The  variations  from  concentration.  32 The  results  indicated  tribution  between  by  application  and  foliar  inhibitory  related  to  levels  the  The  major  was  ethanol  Actigraph  and  whereby  roots  was  of  5 ppm  2,4-D,  of  2,4-D  produced  of  portion  of  acid of  insoluble  scans  pattern  leaves  synthesis  leaves. also  the  a  picric  corporation  of  activity  into  phosphates)  in  the  and  leaf  not  believed  acid  50  P  of  Both  which  appear the  activity  in  these  be  than  to  roots  be and  tissues  RNA. indicated  (nucleotides in  roots  stimulatory  in  to  disthe  compounds  compounds  tissues  in  favor  ppm.  chromatograms  organic stem  in  responses  insoluble  is  balance  upset  insoluble  the  and  but  the  the  more and  insugar  roots. 32  Plants up,  grown  translocated,  phate-free was  in  complete  and  nutrient.  nutrient  retained Also,  less  prior  to  phosphate  incorporation  into  exposure  than  those  soluble  P  grown  , in  organic  took phos-  compounds  suppressed. The  only  radioactive  compound  in  xylem  exudates  from  32 after  to  their  initial  exposure  to  P  was  orthophosphate.  plants  10  hours  TABLE OF CONTENTS  ABSTRACT  ii  LIST OF TABLES  viii  LIST OF FIGURES  xi  LIST OF ABBREVIATIONS  xiv  ACKNOWLEDGEMENTS  xv  INTRODUCTION  1  LITERATURE REVIEW  4  A.  B.  A c t i v i t y of a u x i n in the p l a n t  4  Movement in the p l a n t  4  S p e c i f i c r e a c t i o n s w i t h i n the p l a n t  5  S t r u c t u r e and a c t i v i t y  5  Summary  6  E f f e c t of a u x i n s on m i n e r a l uptake and  incorporation  by p l a n t s Phosphorus Other m i n e r a l s Summary C.  E f f e c t of a u x i n s on growth and c o m p o s i t i o n of the p l a n t Summary  D.  E f f e c t of a u x i n s on c e l l u l a r m e t a b o l i s m  7 7 9 10 10 12 12  R e s p i r a t i o n and p h o s p h o r y l a t i o n  12  Glucose and a c e t a t e u t i l i z a t i o n  14  Enzyme a c t i v i t y  16  The  E.  eel 1 wai 1  16  Photosynthesis  17  Summary  17  Interactions  between  auxins  and p r o t e i n s  or  nucleic  acids  18  Proteins Nucleic  18 acids  19  Summary METHODS A N D  21  MATERIALS of  23  A.  Growth  plants  23  B.  Treatment  with  2k  C.  Exposure  D.  Harvest  E.  Digestion  F.  Chromatography  to P  2,4-D  2k  3 2  and e x t r a c t i o n  25  and c o u n t i n g  26  and s c a n n i n g  27  RESULTS  29  A.  Total  B.  Percentage  distribution  leaf,  and root  C.  uptake  Percentage acid  D.  Percentage leaf,  ethanol  and a c i d  stem of  of  of  the total  activity  among k]  the total  insoluble of  and root the a c i d  soluble  29  tissue  distribution  Distribution the  the plant  distribution  soluble  among E.  stem  by  activity  between  fractions  the a c i d  kk  insoluble  activity  tissue  k8  insoluble activity  and e t h a n o l  insoluble  between  fractions  . . .  51  F.  Percentage among  G.  H.  distribution  leaf,  stem  Distribution  of  of  and root activity  the acid  soluble  activity  tissue within  52 the acid  soluble  fraction  5 6  Leaf  tissue  5 6  Stem  tissue  6 6  Root  tissue  73  Nucleotides  73  Distribution  of  activity  in xylem  exudates  DISCUSSION  80 8 2  A.  Phosphate  uptake  82  B.  Phosphate  translocation  84  C.  Incorporation  D.  Acid  SUMMARY BIBLIOGRAPHY APPENDIX  soluble  into  insoluble  compounds  compounds  88 92 9 5 98 Ill  v i ii  LIST  I.  Volumes  of  stock  OF  solutions  TABLES  used  for various  nutrient  solutions  23 32  II.  Total  uptake  and d i s t r i b u t i o n  plants  sprayed  posure  to P  with  of  P  by c o n t r o l s  5 ppm 2 , 4 - D o n e h o u r  before  and ex-  32 31 32 III.  Total at  uptake  various  and d i s t r i b u t i o n  sampling  of  P  by c o n t r o l  plants  times  32 32  IV.  Total  uptake  and d i s t r i b u t i o n  times  by p l a n t s  Total  uptake  times  by c o n t r o l s  Total  uptake  times  by p l a n t s  sprayed  with  of  P  at  various  sampling  5 ppm 2 , 4 - D  33  32 V.  and d i s t r i b u t i o n and p l a n t s  of  P  at  sprayed  various  with  sampling ..  5 ppm 2 , 4 - D  36  32 VI.  and d i s t r i b u t i o n sprayed  with  of  P  at  various  sampling  50 ppm 2 , 4 - D  38  32 VII.  VIII.  Total  uptake  times  by c o n t r o l s  Percentage stem  and d i s t r i b u t i o n and p l a n t s  distribution  and root  tissue  of  of  of  P  sprayed  the total  control  at  various  with  5 0 ppm 2 , 4 - D  activity  plants  sampling  among  harvested  ..  40  leaf,  at  32 various IX.  times  Percentage stem  after  initial  distribution  and root  tissue  of  of  exposure the total  treated  to P  42  activity  plants  among  harvested  leaf,  at  32 various  times  after  initial  exposure  to P  43  i X  X.  Percentage acid  distribution  soluble  plants  and  harvested  acid at  of  the  total  insoluble  various  activity  fractions  times  after  of  between control  initial  ex-  32 posure XI.  to  P  Percentage acid  45 distribution  soluble  plants  and  harvested  acid at  of  the  total  insoluble  various  activity  fractions  times  after  of  between treated  initial  ex-  32 XII.  posure to P Distribution stem  and  sampling  of  root  the  acid  tissue  times,  of  insoluble control  expressed  as  a  activity  plants  at  percentage  among  leaf,  various of  the  total  activity XIII.  49  Distribution stem after  and  of  root  the  acid  tissue  spraying,  of  insoluble activity treated  expressed  as  a  plants  at  percentage  among  leaf,  various of  times,  the  total  activity XIV.  50  Distribution stem  and  sampling  of  root  the  acid  tissue  of  times,  soluble control  expressed  as  a  activity plants  at  percentage  among  leaf,  various of  the  total  activity XV.  54  Distribution stem after  and  of  root  the  acid  tissue  spraying,  of  soluble treated  expressed  as  a  activity plants  at  percentage  among various of  the  leaf, times total  activity XVI.  Location grams  55 of  standard  developed  solvents  46  in  phosphorus  the  picric  compounds  acid  or  on  chromato-  isobutyric  acid 57  X  XVII.  Distribution and  acid  various XVIII.  acid  spraying XIX.  acid  spraying XX.  sampling  with  with  activity  fractions  the total  between  in control  acid  soluble  plants  at Ill  activity  fractions  at  between  various  acid  times  soluble  after  5 ppm 2 , 4 - D of  the  insoluble  Distribution ethanol  of  total  times  insoluble  Distribution and  the  insoluble  Distribution and  of  total  113 activity  fractions  at  between  various  acid  times  soluble  after  5 0 ppm 2 , 4 - D of  soluble  the acid  115  insoluble  and ethanol  activity  insoluble  between  fractions  the 116  xi  LIST  1.  The e f f e c t  of  at  times  various  5 ppm 2,4-D after  OF  FIGURES  on the t o t a l  uptake  of  P  spraying  34  32 2.  3a.  The e f f e c t  of  at  times  various  Actigraph soluble  3b.  Acid  3c.  Acid  Acid  Acid  sprayed 6a.  Actigraph soluble  6b.  Acid  sprayed  of  P  acid  chromatogram;  tissue leaf  39  from  tissue  acid  control from  plants  . . . .  of  of  leaf of  picric of  leaf  fraction  of  60 leaf  tissue  from  plants 61  acid  chromatogram;  tissue leaf  from  tissue  acid  control from  plants  . . . .  of  stem  fraction  of  5 p p m 2,4-D  62  plants 63  acid  chromatogram;  tissue leaf  from  tissue  acid  control from  plants  . . . .  64  plants  50 ppm 2,4-D picric  59  plants  5 ppm 2,4-D  fraction  with  of  picric  fraction  scan o f  soluble  of  5 ppm 2,4-D  fraction  with  leaf  fraction  scan of  soluble  uptake  spraying  picric of  on t h e t o t a l  5 ppm 2,4-D  scan  with  after  fraction  fraction  Actigraph soluble  5b.  with  soluble  sprayed 5a.  with  Actigraph soluble  4b.  fraction  soluble  sprayed 4a.  scan o f  soluble  sprayed  50 ppm 2,4-D  65 acid  chromatogram;  tissue stem  from  tissue  acid  control from  plants  67  plants 68  xi i  7a.  Actigraph soluble  7b.  Acid  Acid  soluble 9b.  Acid  10a.  Acid  11a.  soluble lib.  Acid  12.  with  Actigraph  of  5 ppm of  chromatogram;  tissue stem  from  tissue  acid  control from  plants  . . . .  root of  70 acid  chromatogram;  tissue stem  from  tissue  acid  control from  plants  . . . .  of  root  fraction  of  72 acid  chromatogram;  tissue root  from  tissue  acid  control from  plants  . . . .  picric of  root  fraction ppm  50  scan  of  of  75 acid  chromatogram;  tissue root  from  tissue  acid  control from  plants  . . . .  77 acid  chromatogram;  tissue root  from  tissue  acid  control from  plants  . . . .  T/R  vs.  79 acid  chromatogram;  xylem 81  hours  to  harvest  for  control  plants  of  two  age  groups 14.  T/R  vs.  sprayed  78  plants  exudate 13-  76  plants  2,4-D  picric  74  plants  5 ppm 2,4-D of  71  plants  2,4-D picric  69  plants  2,4-D  picric of  fraction  with  stem  fraction  scan  soluble  sprayed  of  fraction  Actigraph  picric  ppm  50  of  acid  2,4-D  fraction  scan  soluble  sprayed  5 ppm  scan of  with  stem  fraction  fraction  Actigraph soluble  10b.  with  soluble  sprayed  of  fraction  Actigraph  picric  scan of  soluble  sprayed 9a.  with  Actigraph soluble  8b.  fraction  soluble  sprayed 8a.  scan of  85 hours with  to  harvest  5 ppm  2,4-D  for  plants  of  two  age  groups 86  xi i i  15.  T/Sol. age  16.  hours  to  harvest  for  control  plants  of  two  groups  T/Sol. with  17.  vs.  vs. 5 ppm  Actigraph soluble grown  in  89 hours  to  harvest  for  older  plants  sprayed  2,4-D scan  of  fraction complete  90 picric of  root  acid  chromatogram;  tissue  nutrient  from  control  acid plants 117  x i v  L I ST OF  ABBREVIATIONS  ADP  Adenosine  diphosphate  AMP  Adenosine  monophosphate  ATP  Adenosine  triphosphate  CoA  Coenzyme  est  proportionality  2,4-D  2,4-Dichlorophenoxyacetic  DNA  Deoxyribonucleic  A constant  acid  F-6-P  Fructose-6-phosphate  FDP  Fructose-1,6-diphosphate  Fe  EDTA  Fe+3  Ethylenediamine  G-l-P  Glucose-1-phosphate  G-6-P  Glucose-6-phosphate  GMP  Guanosine  IAA  Indoleacetic  acid  tetra-acetic  monophosphate acid  MCPA  4-Chloro-2-methylphenoxyacetic  NAA  Naphthy1acetic  PGA  Phosphoglyceric  R-5-P  acid acid  Ribose-5-phosphate  RNA  Ribonucleic  acid  RNA-ase  Ribonuclease  s-RNA  Soluble  ribonucleic  UDP  Uridine  diphosphate  UTP  Uridine  triphosphate  acid  acid  acid  XV  ACKNOWLEDGEMENTS  The  author  suggestions research  afforded  reported  his  gratitude  and  during  R.  T.  were  Research  J.  gratefully  thesis.  the most  committee:  Wedding  The  to  h i m by P r o f e s s o r  in t h i s  for  and Dr.  much  like  J.  of  Dr. R.  (University  J.  comments  the manuscript  D.  J.  Wort,  Campbell. of  D.  acknowledge Wort  The a u t h o r  helpful  the preparation  candidate's Tenner,  would  Dr.  a t a l l phases  also wishes  made  to  a s t h e work  by t h e members G.  H.  The s p e c i f i c  California,  the guidance and  N.  Towers,  comments  Riverside)  the  express progressed of  the  Dr.  given  on t h e  of  G. by  M. Dr.  manuscript  appreciated.  r e s e a r c h was s u p p o r t e d Council  of  Canada,  in part  f o r which  by a g r a n t  the author  from  i s most  the  National  grateful.  INTRODUCTION  Thimann promotes  (116)  growth  centrations own  inherent  stated it  that  causes  dole ity  term  "auxin"  view  uses  of  Already this  will  their  Many  systems useful  have  through  of  applied  been  of  The  control  role  that  is s t i l l  hormones  t h e mode  therefore  of  plays  studies  action  of  of  auxins.  to  for  interest growth  More  that  of  and  phenomena  cellular  Much  points  levels  in such  available  thesis.  promise.  the  known.  The  capacity  much  from  is  widespread  stimulatory  not f u l l y  become  their  fundamental  auxin  in t h e i r have  of  in-  various  the most  holds  stem  i s ,  and m o b i l -  in t h i s  from  but  also at  That  is considered.  One o f  crops  further  a s 2 , 4 - D and MCPA,  studied  concerning auxins  new c o n c e p t s  in understanding  sense  has  the d i s t i n c t i o n  compounds  experimentally  and tropisms  which  such  their  occurring  persistence  as h e r b i c i d e s ,  in t h e hormonal  included  these  from  i s an a u x i n .  to another,  which  in low con-  (121)  the n a t u r a l l y  stability,  have  and c o m p o s i t i o n  processes.  like  2,4-D  purposes.  has been  studies  that  i t s general  auxins  applied  Van O v e r b e e k  herbicides,  action in  when  substance  as f a r as p r a c t i c a b l e  one auxin  for different  has been  dominance  biochemists  of  axis  fact  much  and auxin  auxins  the y i e l d  differentiation  from  be u s e d  physiologists  apical  vary  as "an o r g a n i c  substances".  very  discovery,  synthetic  2,4-D  freed  the chemical  t h e mode  purpose.  plant  While  and a p p l i e d  influence  as  reactions  auxins  when  Since  plants  i s an e s t a b l i s h e d  the plant  indole  sharp  of  growth-promoting "It  an a u x i n  the longitudinal  shoots  growth  within  not  along  auxins,...".  between  of  to  has defined  recently, control  may be  further  study  2  is  necessary  however,  to  clarify  such  complex  responses  of  plants  to  auxi ns.  Information  is  needed  whereby  2,4-D  stimulates  present  investigation  concentrations by  the  bean  reactions  of  plant. to  assimilation.  The  tabolism  it  ity on  of  the  the  effects  plants, been  picture  the  placed  of  work on  latory does  and  not  the  imply  that  need  of  alter  to  than  have  (50  other  not  be  the  2,4-D  the  in  growth  little  the  magnitude  the  such  into or  as  account  yield  of  in  of  lower The  nutrition in  the  synthetic  uptake  the  are  the any  (5  use  phosphorus  available  one  intact has  to  the  effect.  being  the  general  in  plants  these  and  me-  emphasis  ppm)  of  and  complex-  metabolism  of  low  cellular  Particular  responses  used,  of  certain  data  the  in which  increases  phosphorus  inhibitory.  factors  taken  Since  actual  on  plays  but  on  way  phosphorus  study  exploratory.  been  ppm)  to  mechanism  Accordingly,  the  attention  phosphorus  to  the  distribution  particular  obtains.  patterns  2,4-D  higher  application 2,4-D  or  plants.  and  focuses  largely  describe  determine  uptake  in  to  intact  to  which  of  used  of  element  levels  rather  of  role  one  been  trends  levels  and  logical  has  growth-regulator  Two  growth,  low  the  approach  which  be  undertaken  central a  can  growth  affect  This  related  makes  the  was  2,4-D  which  stimu-  concentrations time  of  application  beans.  32 The the  distribution  roots,  and  a  of  subsequent  synthetic  processes  site,  incorporation  and  radioactivity  incorporation  associated into  following  with  organic  period,  phosphorus compounds.  brief  uptake  mainly  uptake, Since  of  P  by  represents  movement  to  phosphorus  a is  3  taken  up  and  results little these  is  related  reserve aspects  inhibitory bean  incorporated to  actively  metabolic  phosphorus. of  The  phosphorus  effects  of  low  by  plants,  rates,  distribution  particularly  present  metabolism  the  study are  concentrations  seeks  related  of  2,4-D  in  plants  to  determine  to on  which containing whether  stimulatory the  or  metabolism  of  seedl ings.  The  comparisons  which  have  been  made,  for  whole  plants  or  leaves,  32 stems  and  activity,  roots acid  uble  activity  have  been  some of  cases.  the  organic  insoluble as  made  previously  acid  individually,  grown In  activity  revealed  at  various in  phosphorus  by  and  paper  times  up  to  compounds  has were  the  P  activity,  distribution  70  hours  nutrient,  experiments,  fraction  total  chromatography.  phosphate-free  certain  soluble  include:  been not  the  acid  further  in  soluble  acid  complete  of  plants  nutrient  fraction  subdivided,  sol-  comparisons  treatment  insoluble  identified.  the  These  after  or  of  acid  but  or  in part  individual  4  LITERATURE  A.  Activity  Movement In  in the  t o bean  in the  plant  plant-  leaves  in extracts  bioassay. solution the  auxin  1 9 5 6 Hay a n d T h i m a n n  applied later  of  Nine  of  27 h o u r s  the  occurring  roots  no a c t i v i t y  after  bean  hypocotyl  (50) f o u n d  the lower  hours  to primary  lower  REVIEW  could  hypocotyl  leaves,  Little  and roots  internode  less  drops  using  of  and upper  2 , 4 - D was  a  hours  the s p l i t - p e a  radioactive (68) f o u n d  no r a d i o a c t i v i t y . than  of  up t o 4 8  and Blackman  contained  c o n t a i n e d much  100 u g / p j a n t  be d e t e c t e d  application of  and roots  in the f i r s t  when  the shoots, hypocotyl.  Even  2,4-D  that after  accumulation Crafts  (35) h a s  14 shown a  by a u t o r a d i o g r a p h y  bean  cotyl  plant by f o u r  nificant (l%= to  does  10g/l=  2,4-D In  has  at  made  the  from  into  penetrated  ppm) w a s a p p l i e d than  IAA  the rate  to  1  a source  to bean  a l l stimulated  IAA  when  tops.  petioles  past  than  leaf the  an  of hypo-  insig-  0 . 0 5 % Na-2,4-D  2,4-D  by M c C r e a d y  has been  shown  and Jacobs  (82).  mm/hr. herbicides  is a correlation to a s i n k .  interesting observation  o r down  (62) s t a t e d  the roots  the t r a n s l o c a t i o n of there  to one primary  leaf  Lalova  i n bean of  applied  the opposite  2,4-D  out that  movement  -2,4-D  treatment.  summarizing  food  C  after  slowly  moved  pointed  adenine  of  10,000  move more  The  n o t move  days  amount  that  that  transport  between  Also, ATP,  2,4-D  Libbert  ADP,  through  in plants,  transport  e t a j _ . (66)  AMP, a d e n o s i n e ,  sunflower  (Crafts(35) and have  and  hypocotyls.  They  5  state  that  these  substances  possibly  act  through  effects  on  nucleic  acids.  Specific  reactions  Zenk acid and  which  introduced group  of  the  idea  Structure  and  (55) auxin  physico-chemical valent the  bonding  all  of  2,4-D  CoA.  In  2,4-D  1953  (ll8),  been  found  following  Leopold  complex  in the  involves  indoleacetylaspartic  has a l s o complex  an a u x i n - C o A  an  by  Swets  initial  and Guernsey  involving  the  re-  (65)  sulfhydryl  cell.  hydroxylation  and h y d r o x y l a t i o n  and a c c u m u l a t i o n  as  i n A s p e r g i11 u s n i g e r  has summarized  the  reactions  cellular constituents  attachment  with  substituents  activity  isolating  (44).  activity-  Johnsson mechanism of  in  an a s p a r t y l  is formed  of  in beans  plant-  IAA.  acetyl  that  coenzyme  Detoxication  of  form  requires  the  glucosides  ester to  (114)  the  succeeded  1961  glucose  Wedding  action  in  (138)  and a  within  on  the aryl  compounds,  lation  by  at  a  chemical ring  ring  substituted  phenoxyacetic  chloro  (b)  the aromatic  variously  halogenated  with  isolated mitochondria,  (c)  has been  ideas  reaction  shown  to  regarding as  The be  acids  these  two  lack  herbicidal  a  in c o -  position  (28); and  can uncouple  the  (a)  related  including 2,6-di-and  concentration  :  resulting  chelation.  phenoxyacetic  acids,  sufficient  classical  to  of the  although  2,4,6-triphosphoryaction  and  6  are  not  adsorbed  Cocordano the  electronic  concluded 3 and  that  to  and  s u b c e l l u l a r p a r t i c l e s J_n  Ricard  structure auxin  6 positions  Thimann  with  a  substrate  the  Porter  ortho and  the  position  Thimann  induction  within  favorable  distance  (ll6)  but  (96)  the  and  involved and  to  a  sulfur  out  that  a  atom  position  of  with  substitution  have  interpreted of  a  fractional  carboxyl  group  the  action  ring  in  =NH  at  the  achieve combine  not  the  limited  ring.  terms  charge  in  They  may  is  on  positive  (at  acids.  could  association  auxin  considering  substrate  the  the  by  phenoxyacetic  fixation  that  problem  pointed  varies  the  this  (24).  has  molecule  from  studied  chlorinated  simultaneously  bonding.  to  of  have  activity  such  charged  (33)  vivo  of at  the a  indoles).  Summa r y -  From  these  externally this or  become  occurs  freely  studies  in a  directly highly  throughout  becomes  distributed  to  a  only  probable.  certain  it  is  immediately  apparent  associated with  s e l e c t i v e manner.  the in an  limited  plant  when  applied  asymmetrical number  of  cellular 2,4-D to  the  or  auxins  applied  substituents  does  fashion.  compounds  that  not  foliage  That  move and  auxins  groups  and rapidly soon  can  appears  bond quite  7  B.  Effect  of  auxins  on m i n e r a l  uptake  and  incorporation  by  plants  PhosphorusRoot trations  a p p l i c a t i o n s of of  phosphorus the  sand  noted  the  tent to  and of  be  upward  to  a  50 ml  as  of  to  produce  a 0.1%  plants  treatments  2,4-D  P  K)  (and  weeks  after  ppm)  of  2,4-D  were  noted  a  in  decrease  to  ion  in  controls.  uptake.  phosphorus  14  This days  high  solution  growing,  less  at  adverse  were  two  compared  decrease  translocation  of  much  analyzed  (110)  foliar  shown  tobacco  (1200-9600  plants  or  been  contained  when  Harrison  the  due  young  tops  plants,  have  When  in which  concentrations Smith  auxin  uptake.  that  treated  the  2,4-D  effects was  on  poured  into  W i l d o n _et a j _ .  in  comparison  treatment. sprayed  on  barley  total  The  reduction  treatment  P  spraying,  un-  high plants,  (and  Ca)  was  also  (127)  to  After  the  after  concen-  con-  believed  retarded when  the  32 roots et  of  a l .  the  (128)  were  growing  tops  as  effect On  tion and  of  found at  on  to  other  uptake  that  in  Starchenkov in  immersed 20  ppm  controls under  hand, and  in (122)  P which  the  high-N  some,  lower  wheat  but  found  added  not  that  o c c u r r e d as  to  containing  the  decreased  two  lowest  N  sand the  P  .  Wolf  in which  P content  levels,  but  soybeans of  had  the no  f e r t i l i t y .  incorporation and  solution  levels,  at  much  in a  2,4-D  nitrogen  tomatoes  minerals  increases  that  P uptake  the  were  three  compared  stimulate ported  plants  concentrations in  some  cases.  0.001% 2,4-D all  phytate  promoting and  2,4-D  or  Bryantseva  caused  experiments.  growth  of  In  IAA (l8)  increased corn,  levels  of  may re-  absorp-  Vlasyuk 2,4-D  inorganic phosphate.  caused A  similar tion by  treatment  until  several  Humphries  and  of  and  incorporate  sugar  days  beets  by  these  afterward.  Excised  Maciejewska-Potapczyk more  phosphorus  authors  than  bean  (54)  retarded tops  using  controls  have  5 mg/1  over  P  incorpora-  been  IAA  a period  induced  to  take  of  one  up  week.  32 Fang and leaves,  Butts but  trols,  (4l)  no  when  observed  change  10-100  seedlings growing The p e r c e n t a g e o f  in  a decrease  the  ug o f  stem o r  2,4-D  in  root  was  total  P  activity  tissue,  appl ied  to  in  one  in  comparison primary  the to  leaf  con-  of  bean  f o r 7 days a f t e r treatment in n u t r i e n t c o n t a i n i n g P i n o r g a n i c p h o s p h a t e was h i g h e r i n t h e l e a v e s o f the  32  -4 treated  plants.  A  growth  of  reduce  phosphate  The  e x c i s e d wheat  potatoes, of  fluence and  by  at  organic of  is (10)  have  the  forms  dropped  sharply.  increases  uptake  Williams in a c i d  of  inorganic phosphorus  to  clover.  These  changes  to  may  lead  also  be  and  to  and  forms (40)  stimulated Saddik  the  (112)  to  that  P  phosphorus  can  the  the  sunflower  opposite and  associated with  a  break  under  the  in-  Berezovski  with  50  mg/1  rose  the  have  been  found  effect.  in  the  P  They  concurrent ug o f  50  that  in  fraction.  applications  a p p l i c a t i o n of  up  inorganic  phosphorus  stated  results  taken  spraying  while  of have  phospholipid  small  organic  after  Stenlid  concentrations  that  and  which  controls.  the  reported  soluble,  in  into  P,  by  M)  Faludi-Danie1  phosphorus,  of  (10  inorganic  high  However,  (92)  the and  and  directed  lowered  and  organic  2,4-D  found  of  sufficiently  2,4-D  Ormrod  was  74%  Faludi  forms  2,4-D  Kurochkina  to  between  2,4-D.  2,4-D  roots  uptake  balance  be a l t e r e d  down  concentration of  2,4-D  organi by  observed  decreases per  redistribution  in  plant  the  '9  plant,  Tomizawa  (119)  noted  in  soybeans  that  2,4-D,  at  a  concentration  32 which P  inhibited  from  and  the  p  uptake  leaves  decreased  (98)  have  bean  plants  to  shown  that  caused in  tents  were  higher,  found  increased in  effect  Other  the on  (129)  the  six  in  of  days  and  in  the  following  phosphorus  was  of  acid  later.  inorganic  organs  G-l-P,  total,  unchanged  levels  organic  hundred  to  and  growth  also  Uptake  of  during  the  Rhodes  N,  P  in  FDP.  However roots.  the  to  of  whole  Rebstock  soluble  similar  not  translocation  2,4-D  the  plant,  et  a l .  leaves  of  and  alcohol  soluble  the  stem,  P  con-  e_t a j _ .  (70  in  Loustalot  phosphorus a  found levels  in that in  P,  and  of  2,4-D  the  first  in  treatment  the of  stem  and  beans.  The  determined.  0.002%  K was  whole Cl  same  N  2,4-D  days  after  a  the  Ca,  no  plant  two  weeks were  experiments  both  caused via  was  the  ppm  e_t a]_.  a  decl ine  the  roots  change after  higher  in  and  Wort  roots.  in N  content,  Geranmayeh  and  Ca  unchanged,  treatment  of  MCPA  found  tobacco Mg,  Mn,  than  that  plants and  treatment.  as mentioned  by  by  (127)  of  observed  stems  Mg d e c r e a s e d  roots  and  contents  in  2000  Wildon to  buckwheat  applied  increased,  applied Fe,  on  soluble  tomatoes.  B,  S anc the  and  with  five  on  in  sprayed  i n h i b i t i o n when  (99)  increase  2,4-D  total  faba  first  centration  Na  ppm  increase Vi c ia  and  and  PGA,  in  inorganic  ppm a p p l i c a t i o n  decrease  leaves  other  Spraying  an  a  resulted  increased  into  1000  roots,  minerals-  One  by  a  the  stem,  incorporation  phosphorus  then  the  by  Zn Wolf  controls  above  using  at 20  a  high  resulted and  less  .et i l l all ppm  (48).  conin Cu (128)  nitrogen 2,4-D.  10  Cooke  (34)  number then  of  observed  a marked  minerals  inhibition  for  of  increase over  hours  24  uptake  at  after  times  controls  treatment  in  with  the  100  uptake  ppm  of  a  2,4-D,  and  thereafter.  SummaryPhosphorus tions  of  2,4-D  decreased  uptake and  leading  levels  are  used.  On  most the  to  other  uptake  and  even  at  these  No  consistent  be  suppressed  same  time  plants.with  hand,  by  growth-inhibitory  incorporation  high  manifested  in  inorganic the  stem  growth-stimulatory  incorporation;  low  obtained  the  readily  both  been  at  can  yet  this  into  organic  phosphate tissue  is  not  forms  levels.  when  applications  result  applica-  beans have  always  is These  are increased  obtained  concentrations.  effects  although  on  only  the  uptake  limited  of  other  information  mineral  is  elements  available  in  have  many  cases.  C.  Effect  of  Mi 1 l e r increased was  e_t a_L  by  0.5  inhibitory  that of  auxins  sublethal  numerous  with  10-20  affected  on  (83,  and  84)  1.0  unless  ppm  ppm 2 , 4 - D results.  composition  that  sprays  combined  Wedding  and  found  concentrations  crops.  the  growth  with of  Similar  the  plant  growth  2,4-D  field  beans.  (130,  132)  and  on Wort  can  (126) age  the  juvenile  iron.  2,4-D  e_t a j _ .  although  of  of  be  used  increased fertility  interactions  have  to the of  been  and  yield  Five  has  increase yield the  were ppm  reported the  of  beans  plants  associated  yield  also with  11  foliar  applications  papers  (26,  dicate  21,  33,  a variety  ear  yield,  and  physiological  and  legumes.  total  grain  mersion hours, This  stimulatory  Ball  for  about  noted  14 h o u r s  that  Ross  t h e change  treatment  The ditions They  moisture  x  10"2M  content  by 2 , 4 - D ,  matter after  of  the  authors  of  have  2,4-D.  (102)  spray  the roots  short-term  of  that  content,  on  growth  corn, to  potatoes,  increase  were  on  growth  intermittent  for a period of  im-  of  five  the c o l e o p t i l e s .  in the case of  IAA o r a  The s t i m u l a t i o n  period  to the 2,4-D. showed  levels  of  It  gradlasted  should  a similar  bean  the  stimulated.  effects  elongation  results  to  c a n be a l t e r e d  by t h e r e s u l t s the roots  the moisture  (86)  rates  exposure  beans  ( t o 93% o f  was a p p a r e n t .  beets,  seedl ings  phosphate  and recorded  leaves  treatment,  wilting  Other  status  sugar  in-  leaves  be  pattern after  2,4-D.  a s shown  a p p l i e d 4 ppm 2 , 4 - D  nutritional  initial  hydrolyzable  2.4  Avena  with  improved  observed  (4)  Russian  two y e a r s  sugar  above,  numerous  IAA a n d 2 , 4 - D  only  by a d e p r e s s i o n  and S a l i s b u r y ' s  of  with  of  in the  rate  after  although  a n d Dyke  caused a s t i m u l a t i o n  the normal  of  from  the past  a 0.002% 2 , 4 - D  responses  i n c r e a s e was f o l l o w e d  (eg.  growth)  used  faba,  the long-term  to  effects  in s u n f l o w e r s ,  (48)  Vicia  within  139)  and root  i n 10"^ M I A A o r 2 , 4 - D  return  and  of  Abstracts  (134).  136,  processes  reported.  ual  in  of  111,  Geranmayeh  Besides been  61,  quality  dry weight  have  to potatoes  a small  The stems that  Freiberg  soybeans decrease  controls)  content  reported  of  of  after  and roots hours  certain  and C l a r k  growing  under  con(47). high-N  in the percentage 24 h o u r s .  had dropped  21  under  below  followed after  the  By f o u r  days  controls  t h e same  foliar  dry  pattern.  application  12  of  100  only  ppm  1-2%.  2,4-D a  2,4-D In  applied  maximum  For ponents  25  a  beans,  this to  hours  be  leaves  after  water  of  oils,  content  Bradbury  caused  dipping,  review  vitamins,  may  the  connection,  bean  recent  as  articles  to  the  and  partial  followed  effect  pigments,  of  of  the  Ennis  (17)  stomatal by  increased found  closure  that  by 10  which  ppm  reached  recovery.  auxins  and  tops  on  such  carbohydrates,  diverse Wort's  com(133,134)  consulted.  Summary-  Initial  growth  application on  the  growth  tration, have  for  Effect  and of  or  to  Avery's  influence  and  direct  effect  the  be  the  plant.  cellular  then at  Such  and  response  and  the  followed  nutrition,  applications  on  may  within  of  by  few  hours  long-term  factors  as  environmental the  falls,  low  first  but  plant. the  after  effects  the  concen-  conditions Moisture  change  is  content insig-  concentrations.  metabolism  phosphory1 a t i o n -  (2)  review  in  Bonner  induced of  of  treatment  auxins  respiration.  tissue  occurs  this  application,  after  and  IAA;  composition  foliar  of  Respiration  late  2,4-D  found  shortly  nificant  D.  time  been  rises  of  stimulation  water  the  1951 et  aj_.  uptake  auxin  emphasized  but  (15) and  the  applied  ability IAA  respiration.  related  to  the  act  to The of  of  2,4-D  Jerusalem latter water  to  stimu-  artichoke  was  not  a  accumulation  13  since ham  i t was r e p r e s s e d  (21)  higher  obtained  when  tissue.  of  need of  authors  metabolism,  upon  Switzer harvest, their  (58)  (115)  have  capacity  2,4-D  induced  pointed  growth,  out that  occurs only  in respiration which  i n t h e R.Q.. in the presence often  occur  n o r be t h e r e s u l t  of a direct  effect  2,4-D  18-24  before  of  to soybeans  the chemical  results with  with  the mitochondria RNA.  higher  a r e much  Wedding  responses.  energy  phosphate  Marre  (102)  concluded  retained  than  to 5xl0"^M.  before  increased  more  d i d those  to the  mitochondria  Stenlid of auxins  than  and F o r t i  hours  from  reaction Key e t a l .  isolated  from  harvest.  soy-  During  in nucleotides,  (124, 125)  of  have  phos-  used  in Chlorel1 a and in mitochondria  those  (77)  and p a r t i c u l a r l y ATP  and S a l i s b u r y  down  and Black  plants.  higher  which  directly  2,4-D 24 h o u r s  the concentrations  growth  proposed  growth  u p 0^ a n d p h o s p h a t e  phosphorylation  various  phosphorylation  the  similar  and p o s s i b l y  from  Burling-  and hypocotyl  increases  isolated mitochondria  to take  treated  to uncouple  isolated  5x10"^  Addition  obtained  which were  2,4-D  embryos  observed  inhibition at concentrations  beans  pholipids  t o bean  also  growth,  applied  plants.  caused  have  the increases  and subsequently  untreated  pressure.  i n 0^ c o n s u m p t i o n , b u t CO^ e v o l u t i o n w a s  auxin-induced  with  osmotic  r e s p i r a t i o n (29).  original  flask  (53)  While  n o t be c o n n e c t e d  auxin  Ross  no i n c r e a s e  by 2,4-D.  active  increased external  0 . 5 ppm 2,4-D w a s a p p l i e d  Other  induced  by  and Saddik needed  producing  h a d shown levels  that  their  could  have  to affect typical  oxidative  auxinic  i n 1958 t h a t  high-  be i n c r e a s e d by  results  m e c h a n i s m o f 2,4-D a s a n o x i d a t i v e  (112)  failed  uncoupler,  to  IAA.  support  but were  in  agreement  with  reports  has  from  positive  results,  ie.  ones  differed  with  plant  and  In 10  M 2,4-D  were  for  of  led  ences  pea  at  stimulated which  high  there  energy  the  and  that  inhibited glucose  at  glucose  uptake.  was  the  in  that  this  corn  roots.  a  this  the  a  Likholat  transition to  negative  not  of  concentrations  incorporation; Black  pretreatment  pentose  Bourke of  increased  lead  to  and  any  and  Humphreys  and  observed  or  10  three  (14)  in  2,4-D  fact found  increased  contribution  significant of  glucose  differinto  -3 M.  At  10  M,  concentrations  treated  (53)  (16)  -4  10  all  have  the  in  authors  e_t a j _ .  10~^M  incorporation  of  may  with  pathway  These  treatment  treatment  did  pattern  that  including corn.  concentrations  but  in  reported  c a t a b o l ism v i a  while  higher  (52)  plants  activity  pathway,  controls  Dugger  conclusion  roots  components  2,4-D.  compounds,  -5 tissue  by  involved.  various  activity,  pentose from  to  increased  glucose  of  glycolytic  excised  the  tips  concentration  is  utilization-  increased  later  glycolytic  the  Humphreys  root  increased  the  acetate  1957  excised  that  metabolism  (67)  Glucose  shown  that  etiolated  2,4-D  decreased  corn  seed-  _3 lings  with  10  M 2,4-D  G-6-P  dehydrogenase  phofructokinase,  and  Observations  indicated  that  pentose tissue  pathway. cultures  6-P-gluconate  aldolase  creased.  the  found  and  had  in  intermediates  Scott  enhanced  the  dehydrogenase  increased  recently that  increase  glycera1dehyde-3-P  on m e t a b o l i c  auxin More  an  and  relative  e_t a_L  (106)  of  while  that  dehydrogenase  the  growth  activity  enzyme  activity working  tissues  in  of  phos-  was  de-  studies of  with  the  of  all  the N icotiana  presence  of  IAA was a c c o m p a n i e d 6-P-gluconate while  dehydrogenase  enolase,  unaffected.  by d e c r e a s e d  isocitrate  in a c t i v e l y  way  in older,  only  Bianchetti double  (75)  (76)  creased  t h e amount  glutarate,  Baker  also  shown ratio  found of  tissue.  optimal  exposure  pyruvate  phosphate  a n d Ray  pointed  was  always  But  pretreatment  have  accompanied with  by  increased  10  to decrease  the subsequent  Both  inhibition,  at  (42)  a n d 10  pea,  bean,  10"^M  and corn  of  this  showed  out  glucose  uptake  (113)>  acetate  tissue  respect,  tissue  no  that  of  of  pre-  Marre IAA  pathand  almost  segments.  These  to  IAA  5x10  in-  ot-keto-  while  lAA-induced uptake  cell  glucose  Nance  elongation  in coleoptile  by Humphreys  by 2 , 4 - D  respectively.  were  change.  and s t i m u l a t i o n , uptake  controls,  and the pentose  In t h i s  M 2 , 4 - D was o b s e r v e d  (53)  M (53),  tissue  and a c e t a l d e h y d e  G - l - P and t r i o s e  the  g l y c o l y s i s was more  pea-internode of  of  dehydrogenase  concentrations  in e t i o l a t e d  that  dehydrogenase,  to 50-25%  that  meristematic  that  FDP,  (3)  G-6-P  and malate  concluded  differentiated  t h e TPNH/TPN  authors  they  dividing,  have  of  and t r a n s k e t o l a s e  dehydrogenase  Accordingly,  dominant  activity  and  by e x c i s e d at  been  Dugger  corn  50-150  have  tissue.  roots.  ug/plant  observed  a n d McMahon  (88)  in found  _4 that  2.5x10  s]ices.  They  of  the tissue  it  across Much  acetate it  M IAA c a u s e d  cell of  concluded to hold  a transient  that  the  acetate  influx  IAA a p p e a r e d  of  acetate  into  to enhance  in an exchangeable  form  i n by s t i m u l a t e d  tissue  the  pea  stem  capacity  and t o  transport  membranes. the acetate  since anabolic  may b e r e a d i l y  taken  reactions  oxidized  to  CO  are not always as a  result  of  remains  increased an  as  by a u x i n ,  increased  rate  free or in  catabolic  Enzyme  reactions  53,  88,  113).  activity-  Besides metabolic shown  (42,  to  those  enzymes  pathways,  discussed  the a c t i v i t y  be a f f e c t e d  review  articles  various  plants  Cleland  and Burstrom  by  (131> whose  various  133)  of  in connection numerous  auxin  with  other  treatments  enzymes ( l ,  list  some  fifteen  activities  have  been  shown  summarized  this  situation  (32)  particular  51,  different to  has  been  89).  Wort's  enzymes  be a l t e r e d i n 1961  from by  as  2,4-D.  follows:  The a c t i v i t y o f m o s t e n z y m e s i n v i vo i n c r e a s e s a f t e r application of auxin. I t i s n o t known w h e t h e r t h i s i s a response to a u x i n o r to a u x i n - i n d u c e d growth nor has i t been d e t e r m i n e d w h e t h e r t h e s e i n c r e a s e s r e p r e s e n t i n c r e a s e d a c t i v i t y o f enzyme a l r e a d y p r e s e n t o r s y n t h e s i s o f new e n z y m e . T h e e f f e c t o f a u x i n _m v ? t r o o n e n z y m e a c t i v i t y i s e i t h e r i n c o n s i s t e n t between t i s s u e s o r negligible.  Freed  e_t a j _ .  physically ability.  systems appear  The  adsorbed The  standing  (45) to  have  t h e enzyme  contributions  of  enzyme  have  not  presented  of  regulation,  as yet  been  a hypothesis  protein  Monod  thus  whereby  altering  and o t h e r s  (86,  allosteric proteins  applied  to auxin  2,4-D  its  123)  to  catalytic the  and c e l l u l a r  effects,  is  although  undercontrol they  applicable.  e e l 1 wa11 -  Leopold cell-wall  (63)  has concluded  plasticization,  after  summarizing  the synthesis  of  earlier  new w a l l  works  material,  that  and  osmot  uptake  of  water  enlargement stimulate tissue cell  thus  is  an  cyanide  increase  but  not  Actinomycin  lation  to  increased  of  large  (3)  into  the  cell  synthesis, by  both  cell  have  and  which  of  is  constituents and  methyl e s t e r a s e have  cell  IAA  can  coleoptile  glucose  recently  has  deformabi1ity  that  indirectly,  since  uptake. fractions  reported  completely  antibiotic  (87)  elongation  found  upon  polysaccharide  has  (31)  auxins  wall  in a l l  This  Morre  cell  pectin  auxin-induced  D.  of  increased  increases  in e x t e n s i b i l i t y  tissue  action  Ray  Cleland  sensitive  of  and  accompanied  Similarily,  2,4-D-induced  roles  glucose  by A c t i n o m y c i n  sets  The  Baker  elongation.  elongation.  and  by  found  (97)  lAA-induced  causes  of  always  induced  IAA  b a s i c mechanisms  growth.  directly,  authors  of  the  incorporation  both  during  by  and  elongation  These  IAA  are  that  inhibited  could  block  lAA-  recently  implicated  (RNA  protein)  in  in  re-  and  two  growth.  and  (73) been  Ca  (20)  questioned.  Photosynthes i s -  While of  depression  2,4-D a t 0.01%  little and  the  or  no  Bidwell  occurred  (46)  direct (l20)  and  photosynthetic  0.001% (72)  evidence  found  following  of  that  is  available  pulses  application  to  of  rate  beans  following has  been  regarding  of  c'^O^  10-50  ppm  uptake sprays  the  application  demonstrated,  stimulation.  Turner  of  duration  of  an IAA  hour's to  bean  leaves.  Summary-  Some  of  the  factors  which  have  been  found  to  be  important  in  under-  18  standing  the  duration, of  the  responses  of  mode o f  the  and  plant  Reactions  tested  which  and  are  plant  treatment;  the  at  a  lower  one.  above  have  been  shown  in  pation  of  point the of  in  auxin  which  growth they  (108)  has  observed  E.  and  may  are  the  by  concluded  probably  Interactions  to in  be  the  that  result  (64)  phenomena  auxins  and  of  of  the  or  auxins  is  they  may  at  particia  particular  that  the  one  of  diversity  influence,  to  nucleic  and  Setterfield auxins  fundamental  or  be  mentioned  action.  more  part  effects.  direct  responses  proteins  and  indirect  concluded  unitary  results  and  which  metabolic  nutrition,  reaction  of  concentration,  responses  has  action  the  concentration  the  a particular  the  age,  one  of  underlying  are:  direct  at  any,  Leopold  secondary  between  of  if  hormonal  an  auxins  species,  auxins  differentiation  mediate  also  vivo  pathway.  c h a r a c t e r i s t i c s of the  by  Few,  molecule  a metabolic  to  separation  inhibited  stimulated  the  tissue  usually  effects.  acids  Prote insSix 2,4-D of (93)  days  to  the  one  stems  have  after of  the  was  been  of  Recently  and  promote  fresh  incorporation  primary  double  able  incorporation Fang  application  to  amino Yu  weight into  of  retard  (43)  of  the  drop bean  into  increases  whereas  1000  using in  ppm  the  (107).  that  excised  a  plants,  by  proteins  reported of  of  controls  senescence  have  protein,  one  leaves  that  acids  of  2,4-D  pea  inhibitory  at  protein  Osborne  detached IAA  solution  to  content  and  Hallaway  maintain  leaves  of  levels  enhanced decreased  the  Prunus.  concentrations  shoots,  of  which  glycine glycine  19  absorption  and thus  centration  of  was found  (90).  Cleland  coleoptile  Knypl by  also  with  retards  protein  pea  to  sections  Nucleic  in  M 2,4-D  protein  proline.  the growth  incorporation  conditions  Thimann  into  synthesis  of  growth  synthesis.  and 2,4-D-induced  IAA  et a j _ .  Setterfield resulted  (57) f o u n d  resulted  the protein/RNA  (78)  which  could  of  the growth have  fractions  48  spraying  growth  induced  The  the  same  nuclear  was n o t  found  of  pea  green  De H e r t o g h  et  to  substance.  cells  in areas  to  cultures  D but  al.(8).  by B a s l e r  that  decrease  division.  auxin  expansion.  part  RNA at of  in the  and o t h e r s  with  a n d RNA w i t h  restored  localized  and a  cell  nuclear  The p r i m a r y be  of  nucleotides  stimulate  seedlings  content,  hormone-induced  by A c t i n o m y c i n  shown  soybean  t h e RNA  hours  in a r t i c h o k e during  been of  of  soluble  e_t a_K ( l 02)  be a b o l i s h e d  synthesis  microsomal  increased  synthesis  by Roychoudhury  centration  within  (108) o b s e r v e d  i n RNA  that  in a doubling  ratio  shown  RNA  for  inhibited  Protein  and  auxin-induced by  con-  NAA-induced  by Nooden  inhibit  has  and  2,4-D  was r e v e r s e d  acid  (37)-  the experimental  and Hanson  Mashtakov  on  amino  IAA,  t h e same  acids-  Key 5x10  however,  ine to  inhibited  segments  specifically,  synthesis  sections  also  lAA-induced  under  protein  inhibition  stem  which  internode  be n e c e s s a r y ,  stem  This  More  inhibited  hydroxyprol  sunflower  puromycin,  which  inhibit  sections.  antibiotic of  to  (30) u s e d  (60) u s i n g  IAA  incorporation.  chloramphenicol  growth  in  its  and  2,4-D. kinetin  IAA  has  been  synthesis a higher  these soluble  (5, 7» 25).  con-  effects and  20  Increases synthesis creased  since  induced  RNA-ase  Shannon's  by l o w l e v e l s  activity  by g r o w t h - p r o m o t i n g  levels. spect  i n RNA  of  supported  RNA-ase.  that  IAA c o u l d  only  acid  synthesis  b u t n o t de novo  Roychoudhury  stimulate  in both  in  excised  of  auxin  P  and C  tissues  concentration  absorption  related  and Hansen  vented  cotton  excess  of  2,4-D  of  by Sawamura  ration  acid  can induce  has been  inhibitory  conclusion with  reactions  purine  re-  related  i n 1959 to nucleic  and p y r i m i d i n e  (59) h a v e  root  incorporation These  of  bases.  observed  10  these  M 2,4-D, fractions  i n -  RNA a n d DNA  detected That  an  2,4-D  (94) w h o s u p p r e s s e d with  and s o l u b l e but at  t h e same  induced  by  of orientation treated  with  as the basis  2x10  effect c a n be  phosphate M 2,4-D.  RNA s y n t h e s i s time  pre-  t o RNA.  of  For  metaphase  25 ppm 2,4-D. of  of  chromosomes  nuclear  divisions  demonstrated  abnormalities  involving  and m u l t i n u c l e a t e in onion  An  the abnormalities.  2,4-D a n d s i m i l a r h e r b i c i d e s d o w n t o 0.001% h a v e (105) t o p r o d u c e  in  sugars.  are not restricted  were  was p o s t u l a t e d  authors  in pea e p i c o t y l s  the lack tips  into  incorporation.  microsomal  interactions  A l 1 i um  a n d movement  RNA  with  RNA f r o m  nucleic acid  auxin.  synthesis  (36) o b s e r v e d  when  Concentrations used  of  and Blackman  (6) i n h i b i t e d  of  Croker  chromosomes  by P a l m e r  t o RNA  Auxin-nucleic example,  with  on the rate  cotyledons  the loss  -nucleotide  treated  was noted  excised  synthesis  at  a n d S e n (13) s u g g e s t e d  phosphorylation  net  14  inhibitory  Basler  a similar  Biswas  represent  b y K e y (56) t o b e i n -  and reduced  a n d S e n (103) a n d K e y a n d S h a n n o n 32  creases  2,4-D m u s t  found  concentrations  (109) d a t a  to the activity  has been  of  cells.  impeded Also,  been sepa30 ppm  r o o t s (95).  as a requirement  for auxin-induced  growth.  21  Thus,  Masuda  cultures 137)  by  have  (79)  pretreatment  also  functional  in  found  the of  a  cell  wall.  terferes which ment  quired  two  thus  a  with  pea  hour  basis  of  and  cytidylate  to  the  sections with response  time  the  using  s-RNA  stem  after  period  activity  these has  IAA.  been The  and  tissue  RNA  RNA-ase,  C,  RNA  or  two 2,4-D  resulted mainly  its  formation  was  presence or  continued  synthesis  by in  reoccurred,  auxin-s-RNA  et  al.(8,9)  resistance  adenylate by  The  response.  the  increased  inhibited  com-  auxin  Bendana  with  Cp  (38).  growth  in  in-  pretreat-  hours  with  lie  with  Actinomycin  in  may  re-  which  inhibition  the  was  growth  Similarily,  IAA  81,  was  has  (90  by  D for  demonstrated  was  of  growth  interactions  association  (80,  lAA-induced  reduced  complete  kind  Mitomycin  to  artichoke  associated  pretreatment  preformed  recently  RNA  while by  on  extracts  this  synthesis.  hour  before of  that  of  Oota  Actinomycin  the  one  auxin-RNA  pancreatic  regions,  of  and  greatly  RNA  2,4-D  Yanagishima  yeast.  affected was  and  involving  reported  DNA-directed  inhibitor  which  the  has  and  of  layer  expansion;  Masuda  it  association  of  by  replication,  the  2,4-D  (60)  phenol  Avena,  DNA  indicating  The  presented  effect  Masuda  the  tissue  not  nullified of  from  s e c t i o n s was  with  addition  RNA  stem  green  pletely  RNA-ase.  artichoke,  Knypl  interferes of  stimulatory  with  that  among  hypothesis  sunflower  the  auxin-induced  cross-reacting viewed  reduced  and  Actinomycin  D.  Summary-  Either RNA  appears  Protein  the  necessary  synthesis  for  is also  the  occurrence of  necessary  in most  IAA-  of  a  and  cases.  certain  kind  2,4-D-induced Both  of growth.  nucleic acid  and  protein and of  synthesis  inhibited auxin  by  action  are  generally  inhibitory in mediating  stimulated  levels these  of  the  during  auxin-induced  growth-regulators.  reactions  may  be  s-RNA.'  growth  The  site  METHODS AND  A.  Growth  Bean moist in  of  flats  plants  seeds  paper  (Phaseolus  towels  of  vulgaris  f o r 3-4  vermiculite  days.  conditions  66% r e l a t i v e  humidity,  18-20°C,  70%  humidity.  Experiment  Table  I,  V)  60  liters  nutrient  Volumes  of  of  room  Solution  MgSO^  Ca(N0 ) 3  KH P0^ 2  KC1 Fe  EDTA  Micro  2  were  used  cycle:  -P  2.3  .55  2.3  4.5  1.10  2.3  .55 —  for In  nutrient  Strength  —  16 8  transferred  wa t e r  1/4  nutrient  day c y c l e :  aerated.  for various  planted  complete  night then  between  were  (or complete  and the s o l u t i o n  solutions  Complete  candles;  The p l a n t s  m 1 /1  Stock  were:  placed  seeds  strength,  a phosphate-free  solution,  stock  1/4  800 f o o t  were  sprouted  Growth  22-25°C,  containing  Uniformly  Crop)  with  f o r 5-6 d a y s .  relative  v a r . Top  and watered  solution  tank  MATERIALS  4.5  2.3  1.0  .25  1.0  1.0  .25  1.0  hours,  hours, to a  5 ppm most  solutions.  24  experiments I,  IV,  and  The (Table 1.0  one V)  sample  held  in a  nutrient  I)  of  2.86  g,  H_0  .025  B.  Treatment  g>  following  1.0  MnCl2.4  M KCl, HO  all  1.81  with  50 mg o f  one  liter  of  water.  the  primary  prevent until  sections  leaves  of  from  the  plants  solution when  were  the  not  Following for  the  C.  Exposure The  to  were  made  by  using  the  Experiment  appropriate  Fe  EDTA  (5 m g / m l ) ,  and m i c r o n u t r i e n t s  g,  ZnCl2  in one  the  .11  g,  liter  tank of  and  the  plants  the  solutions plants  dripped  of  nutrient  5 ppm  cork.  M MgSO^,  CuCl2.2  of  h^O  1.0  volumes  M  C a t N O ^ , (HjBO  . l l g ,  water).  2,4-dichiorophenoxyacetic  spraying,  P  in  1.0  the  with  dissolved  in  atomizer  to  an  14  days  placed  in  a bottle  leaves. so  was  were  solution.  returned  plants  acid  applied  they  roots  nutrient  nutrient  the  were  when  the  off  were  Tin common  duration  (three  solutions:  These  contamination  the  plants  2,4-D  or  removed  two  stock  dissolved  Five  were  single  solutions  the  M KH2P0z+,  c o n s i s t e d of  Spraying  The that  old.  tank the  was  The of  was  water  and  to  continued  divided  treated  plants  into  control  solution.  were  kept  in  continuous  illumination  experiment.  32  solution  to  which  the  radioactive  phosphate  was  added  32 was  a  complete  solution,  except  where  noted.  The  P  (30-50  uc/1)  was  25  added  as  quired the  orthophosphate  to  the  total  for  each  experiment.  At  plants  were  transferred  from  holding  one  Experiment  sample,  or  I only),  to  a  which  volume  various the  plastic  growth  the  radioactive  solution  was  replaced  of  aeration  and  illumination  samples;  times  tray  contained  of  after  exposure after  solution  treatment  with  tank  to  individual  holding  up  to  12  nutrient.  exposure  each  were  similar  2,4-D  bottles  samples  radioactive of  re-  that  1 or  2  ppm  The  sample  to  (5  or  group  during  the  32 growth is  period.  referred  ferred with 2-3 to  to  an  The  to  a  as  root  their  pending  at  the  drain  the  made and  rate  respective  to  P  "exposure  washer  overflow  minutes  exposure  of  parts  on  the  experiment.  Harvest  and  Extraction  continued  time".  from  The  a metal  a water  inlet  ca.  1/min.  of  3-5  the  tank  This  plants  were  container hose.  is  (ca.  samples  held  for  hours, then  Washing  The  and  period  for  2,  referred  trans-  700  ml)  fitted  continued were  8 or  to  which  as  for  returned 9 hours  the  de-  "holding  t ime".  D.  At tank,  the  the  roots,  roots  stems  expressed were  on  taken.  cedures  For was  end  the  dried  plus  in  done  a  between  plant  extraction in  extraction,  placed  holding  petioles,  a whole If  were  of  cold  a  paper  and  was  mortar,  to  room  plant  the  (5 be at  and  were the  Except  carried  removed  plants  where  ppm E x p e r i m e n t out,  the  dissected  the  results  I only), this  from  and  fresh the  into  were weights  above  pro-  5°C.  material  frozen  plants  towels  leaves.  basis  cold  the  time,  in  from liquid  each of  the  nitrogen,  three and  organs  ground.  Ten  26  or  15 m l  added  on the weight  and the e x t r a c t i o n  minutes, washing ture  (depending  after  which  continued,  the contents  ( 1 0 - 1 5 ml w a t e r )  was n e u t r a l i z e d ^  with  15 m i n u t e s .  which,  after  centrifugation,  acid  insoluble  In  30 m i n u t e s .  After  supernates  were  fraction  soluble  fraction.  Digestion  The  and  fresh  to  fraction,  N HCIO^ was  grinding,  were  tube.  with  5-10  the previous  mix-  at  ml o f  water  supernate  residue  10-15  with  The  and c e n t r i f u g e d  t h e washed  for  transferred  centrifuge  was washed  was added  0.3  intermittent  the mortar  residue  soluble  IV,  this  t o pH 3 . 0 w i t h  cooling  ml e t h a n o l - e t h e r  soluble  E.  The  t h e 5 ppm E x p e r i m e n t adjusted  The  of  of  thus  being  the  fraction.  80% e t h a n o l ,  20  with  10 N K0H t o p H 7 - 8 ,  g for  the acid  the sample)  t o a 50 m l p l a s t i c  27,000  constituting  of  (3:1)  residue  60% HCIO^,  and c e n t r i f u g i n g , was made a t  collected  and the f i n a l  was e x t r a c t e d  room  a second  the a c i d  the acid  20 m l  a t 45°C  extraction  temperature  and c o n s t i t u t e d residue  by h e a t i n g  with  f o r 30  for  with  minutes.  insoluble,ethanol  insoluble,  ethanol  in-  Counting  plant  material  and i n s o l u b l e  residues  were  placed  in  1 N e u t r a l i z a t i o n j u s t p r i o r to c e n t r i f u g a t i o n decreased the time d u r i n g which a c i d h y d r o l y s i s o f phosphate e s t e r s c o u l d o c c u r and o b v i a t e d c e n t r i f u g a t i o n t o remove t h e KCIO^. The p r o c e d u r e d i d not i n troduce any a c i d i n s o l u b l e r a d i o a c t i v i t y into the s o l u b l e fraction.This was i n d i c a t e d by t h e a b s e n c e o f a c t i v i t y a t t h e o r i g i n o f p i c r i c a c i d chromatograms o f t i s s u e e x t r a c t s in which the percent o f the t o t a l a c t i v i t y i n a c i d s o l u b l e , o r g a n i c compounds (but n o t i n s o l u b l e compounds) was v e r y s m a l l ( F i g u r e 17, Appendix).  27  100 to  ml d i g e s t i o n each  one.  flasks,  The f l a s k  then  3 m l 70% HNO^ a n d 2 m l 6 0 % H C I O ^ w e r e  was heated  until  a l l the plant  added  material d i s -  solved.  The digests ml  acid were  of these  Century nected  solutions  to the scaler  After  on  ml  Whatman  No.  transferred each  spot  papers  course with  of  the acid  (31  : water  the radioactive  Standard  Model  and the  f l a s k s . Ten  G-M c o u n t e r  tube  (20th  in a lead  castle  con-  151A).  and decay  fractions  a t 30-32°C.  The average  when  o f two  necessary.  cm x 21  cm) u n t i l  later.  After  scanner  chromatograms  were  1  These  were  run using  acid,  R-5-Pj  pyrophosphate  (P-O"*1'),  FDP,  phosphocholine,  in a  (69) f o r 2 0 - 2 4 strips  model  spotted  placed on  run in a p i c r i c  scanned  to  a c t i v i t y was  solution  glass acid:  hours.  along the  a t 3/4 in/min  1025).  the following  A D P , A T P , U D P , U T P , GMP, F - 6 - P ,  phytic  were  f o r 1 hour  inch wide  (Nuclear-Chicago,  were  of  equilibration  and c u t into  concentrated  sufficient  The amount  t h e chromatograms  material.  were  The c o n c e n t r a t e s  (2 g : 8 0 m l : 20 m l ) s o l v e n t  a i r dried  II  soluble  detection  chamber,  an A c t i g r a p h  supernates,  t o 5 0 ml v o l u m e t r i c  a liquid  f o r background  was n o t c o n s t a n t .  were  into  soluble  M 6 ) , which was housed  evaporator  to allow  tert-butanol  washing,  (Nuclear-Chicago,  1 paper  chromatographic  The  type  ethanol  and Scanning  counting,  in a flash  with  was poured  was c o r r e c t e d  Chromatography  2-3  supernates,  transferred,  Electronics,  countings  F.  soluble  G-l-P,  compounds: G-6-P,  and inorganic  PGA,  orthophosphate.  28  These  compounds  e_t a j _ . light  (22), until  In the  one  the used  were  heating the  5 ppm for  one  with  to  VI,  and  the  spotted  chromatogram  was  developed  standards  ml:  given  located  by  33 m l : above  molybdate 85°C  1 ml) were  scanning  additional  first  and  (66  at  from  water  ammonia  the  and  reagent  exposure  of to  Burrows ultraviolet  developed.  scanning,  eluted  using minute  Experiment  corresponding  was  for  cojor  region  This  revealed  on  this  major a  in  solvent  used  strips  with as  part  of  area  second an  spots  of  this  the  system  described  acid:  hours. and  above.  beside  chromatogram  chromatogram  12-14  placed  radioactivity  isobutyric for  were  (15  The the  was  cm x  water:  the  27  cm).  25%  first  five  radioactivity  29  RESULTS  A.  Total  uptake  by  the  plant  32 Plants  exposed  orthophosphate  for  to a  a  nutrient  period  counts/minute/gram  fresh  ure  uptake  of  the  time. and  This  there  These  the  was  as  no  allowed 124%  (Table level  was  of  as  uptake,  in  the  met  hours  here by  by  provided  loss  of  the the  plants  presence  up  plant  root  was  and  taken  during  washing  during  grown of  P  the  in  only  carrier  radioactivity.  roots  activity  for  the  took  whole  phosphate  valid  indicated  (5  3 plants, stand  much  for  activity  This  as  a  this was  The meas-  exposure  complete  holding  period.  phosphate-free  traces  of  activity  ppm  Experiment  one-half 1  of  hour,  as  the  difference  I)  which  the  were  treated  controls  was  involving  sprayed  plants  during  statistically  16 s a m p l e s ,  a  1  with  each  5 ppm  were  found  hour  exposure  s i g n i f i c a n t at  2,4-D  to  take period  the  2.5%  7.67).  (F^=  To  2  containing  solutions.  to  II).  of  significant were  1 or  weight  is  an e x p e r i m e n t  consisting  up  assumption  holding  In  and  of  requirements  nutrient, in  rate  of  solution  investigate  the  a  experiments  series  of  effect  of  time  was  after  set  up  spraying including  on  the  this  rate  of  variable.  The  32 uptake Table than  P  111. in  justed time  of  the so  zero.  by  the  controls  in  three  Since  the  exposure  other  two  experiments,  that  all  three  time  in for  experiments (The  such experiments the  5 ppm  next  page  given  Experiment  comparison  approached  is  the  the  values  same  i s 31.)  III  in was  were  total  less  ad-  uptake  at  Table  II  sprayed ment  Total with  5  uptake  and  ppm 2 , 4 - D  d i s t r i b u t i o n of  one  hour before  P  32  by  e x p o s u r e, t o  Control  Average Treated  ( 5 ppm  Experi-  (cpm*)  Root  Stem  519 740  262 371  2,821  637 607  281  2,080  267  598  314  111 656  341 448  1,925 1 ,883 2,651 2,202  673  424  2,249  2,795 3,769 3,306 3,346  651  338  2,183  3,172  730  262 338  2,180 3,301 2,440  3,172 4,586  Total  1,654  233  2,435 3,932 2,998 2,799  3,545 4,011  1 ,065  314  2,632  705 979 806 678  353 444 444  3,813 4,553  262  2,755 3,130 3,181 2,318  Average  848  332  2,743  3,923  Control  130  98  125  124  Counts/minute/3 plants. Exposure time 1 hour. Holding 9  plants  Leaf  947 872  *  P32  and  1).  R a d i o a c t i v i ty'  % of  controls  hours.  time  (time  from  exposure  4,431 3,258  to  harvest)  32  Table  III.  various  Expt.  Total  sampling  uptake  and d i s t r i b u t i o n  P  32  by c o n t r o l  plants  at  times*.  Radioactivity  and  Sampling  of  (cpm/g**)  Times  (hrs.)  5 ppm Expt.  Root  Leaf  Stem  2,530  2,155 2,000  10,060  2,680  16,250 13,900  Total  II  2,695 3,980 4,730  14  2,465 2,580  60  14,745 15,045  10,350  22,910 21,095  13,550  3,335  16,750 12,150  29,035  0  290  258  984  1 ,532  16  394  225  1 ,230  1 ,849  60  655  440  1,520  2,615  5 ppm Expt.  I II  50 ppm Expt. I 4,670 6,340  1,374 1 ,508  10,470  16,514  20  5,260 8,920  2,770 3,450  15,990 13,680  24,020 26,050  48  10,580 9,080  3,760 3,610  16,300 15,350  30,640 28,040  "  Sampling  times  correspond  * *  exper iment. Counts/minute/gram  fresh  Exposure time: 2 hours 1 h o u r f o r 5 ppm E x p t .  to hours  after  spraying  plants  in each  weight.  f o r 5 ppm E x p t . I I a n d 5 0 ppm E x p t . III. Holding time: 2 hours f o r a l l  I; experiments.  Table  IV.  times  by p l a n t s  Expt.  and  Sampling  Total  uptake sprayed  and d i s t r i b u t i o n of P with  at various  sampling  5 ppm 2 , 4 - D .  Radioactivity  (cpm/g*)  Times  (hrs.) Leaf  Stem  Root  Total  5 ppm Expt.  II 2,960  2,720  13,660  19,340  4,345  2,560  12,930  19,835  5,380 6,650  2,695  7,865 9,500  15,940  2,180  14  3,490 5,270  2,275 2,425  15,600 13,850  21,365 21,545  30  6,950 9,160  3,890 3,445  13,750 12,250  24,590  2,720  2,730 4,500  14,750  20,200  7,940  16,650  29,090  2  460  310  1,320  2,090  4  490  360  1,350  2,200  8  225  240  910  1 ,375  16  190  240  1 ,430  1 ,860  30  590  470  1 ,210  2,270  60  160  320  1 ,300  1 ,780  2  6  60  i ppm :xPt.  *  18,330  24,855  in  Counts/minute/gram fresh weight. Exposure t i m e : 2 hours f o r Expt. II a n d 1 h o u r Holding t i m e : 2 hours f o r both experiments.  f o r Expt.  III.  Figure  1.  The  effect  of  5  ppm  2,4-D  on  the  total  uptake  of  at  various  times  after  spraying.  10  35  These  values  results were  and  obtained  plotted  uptake  were  in  plotted  for  the  this  way,  which occurred a  that  nearly  after  the  between  about  same o r  the  creases'  in  were  grown  under  controls  not  present  at  70  the  treated  (Table uptake  nutrition  hours  70  prior  to  retained  about  one-fifth  in  -P  solution. plants  Also, grown  during  in  the  took  increase as  up  Even  nutrient  holding lost  at  aged.  in-  treatment plants  difference  V).  These  as  those  period,  about  phosphate  although  spraying.  as much p h o s p h a t e 8 hour  plants  after  no  in  variation  hours  showed  IV)  phosphate  that  the  maintained;  The  the  indicated  after  (Table  not  controls.  few  When  increases  (5 ppm E x p e r i m e n t  the  complete  the  plants  treatment  spraying  and  that  also  1).  5 ppm 2 , 4 - D  occurred a  after  up  to  V)  (Figure  s p r a y i n g were  than  hours  took  untreated  after  rate  cpm/g  with  hours  and  50  of  apparent  samples appeared  phosphate  full  terms  sprayed  was  slower  experiments  total  these  it  few  a  in  plants  6 hours  at  treated  Subsequent  a  then  from  plants grown  treated  10% o f  the  and total  32 P  which  they  Next,  the  vestigated. Initial until  These  hours  difference  the  up. to  the  results are in  the  after  between  and  taken  response  increases  3-4  spraying  had  the  same  rate  those  given of  and  slightly  concentration in  Tables  phosphate  found  treated or  higher  at  the  control less  VI' a n d  uptake  5 ppm  2,4-D VII  were  level.  plants  uptake  of  at  at  was  and  not  times  Figure  2.  observed  There 20  in-  hours  was  little  after  thereafter.  36  Table by  V.  Total  controls  and  uptake  and  plants  sprayed  E x p t . and Sample  d i s t r i b u t i o n of with  5 ppm  P  32  at  various  sampling  2,4-D.  Radioactivity  (cpm/g* or  %)  Leaf  Stem  Root  Total  6,560  2,520  5,330  14,410  6,960  2,510  5,060  14,530  7,100 4,960  2,890 2,890  5,140 6,070  15,130 13,920  89  115  109  100  Control  5,415 8,515  2,290 2,600  4,920 4,840  12,625 15,955  Treated  6,470 6,040  2,770 2,995  5,290 5,420  14,530 14,455  90  118  110  100  5 ppm E x p t .  50  IV  hr.  Control  Treated  % of  70  Control hr.  % of 5 ppm  50  Control  Expt. hr.  V**  Control Treated  % of  70  Control  840  410  2,150  3,400  415  240  1,385  2,040  885 360  470  270  2,355 1,510  3,710 2,140  101  114  109  108  740  455  2,000  3,195  600  390  1,965  2,955  845  570  2,135  3,550  385  415  1,825  2,625  92  116  100  100  hr.  Control Treated % of  Control  times  37  Table  V.  (cont.)  Expt.  and  Radioactivity  (cpm/g* or  %)  Samp 1e Leaf  5 ppm E x p t .  Stem  Root  Total  VI  1 hr. Control  10,480  Treated  % of  * *  11 , 3 3 0  11,760  25,525 28,760  11,950  5,870  13,200  31,020  11,980  5,860  10,880  28,720  110  115  108  110  Control  Counts/minute/gram fresh weight. Exposure time 2 hours, h o l d i n g time P l a n t s grown i n c o m p l e t e n u t r i e n t .  It  was n o t e d  inhibited  by  the  Similar are  not  in  terms  three  that  considered of  the  organs.  leaf  50 ppm  data  for  10,500  4,545 5,670  for  all  experiments  and stem e l o n g a t i o n  were  somewhat  treatment.  the  here  expansion  8:hours;  individual  on a cpm/g  percentage  organs  basis,  d i s t r i b u t i o n of  are given  as  they  the  in  the  are better  total  activity  tables  but  presented among  the  Table  VI.  times  by p l a n t s  Sampling  Total  uptake sprayed  and d i s t r i b u t i o n o f P with  Times  50 ppm 2 , 4 - D  at various  (50 ppm E x p e r i m e n t  Radioactivity  sampling I).  (cpm/g*)  (hrs.) Leaf  Stem  Root  Total  3,510 4,580  1,030 1,440  10,830  16,850  6,680  1,965 1,940  16,030  5,960  15,970  24,675 23,870  20  8,090 6,560  2,800 2,300  14,290 12,810  25,180 21,670  48  11,260  4,155 4,375  13,240  28,655 26,235  3  9  8,510  "  Counts/minute/gram fresh weight. Exposure time 2 hours, h o l d i n g time  13,350  2  hours.  Hours  after  Spraying  vo  40  Table  VII.  Total  times  by c o n t r o l s  (50 ppm E x p e r i m e n t  uptake  and d i s t r i b u t i o n  and p l a n t s  sprayed  with  of  P  at  50 ppm  various  sampling  2,4-D  II).  Radioactivity  (cpm/g*  o r %)  Sample  10  Treated  % of  Control  Root  Total  6,170 5,005  2,575 2,285  7,090 6,380  15,835 13,670  5,445 3,620  2,780  6,060  15,185  2,875  6,360  12,855  81  116  99  95  5,685  2,490  5,830  14,005  6,435  2,550  5,835  14,820  4,695 7,660  2,810 2,700  6,855 5,800  14,360 16,160  102  104  109  106  3,220 3,650  2,160  8,160  13,540  2,660  7,500  13,815  3,060 4,600  1,590 2,070  6,680 7,660  11,330 14,330  111  76  hr. Control Treated  % of 70  Stem  hr. Control  20  Leaf  Control  hr. Control Treated % of  Control  Counts/minute/gram fresh weight. Exposure time 2 hours. Holding time: s a m p l e s , 2 h o u r s f o r 70 h o u r s a m p l e s .  92  8 hours  for  94  10  a n d 20  hour  41  B. root  Percentage d i s t r i b u t i o n of the total a c t i v i t y among leaf,  stem and  tissue If bean plants grown in -P nutrient were exposed for 1 hour to a  32 nutrient solution containing P  of high s p e c i f i c a c t i v i t y without  c a r r i e r phosphate, and then held for up to 48 hours, 95% of the total a c t i v i t y remained in the roots and about 1% reached the leaves.  However  32 if the P  was contained in a complete nutrient s o l u t i o n , the  transfer  of a c t i v i t y to the stem and leaves was increased (Table VI I I) to 35-65% depending on the holding time. 32 The roots of control plants exposed to P  for 1 or 2 hours and  held for 2 hours contained about 55-65% of the total a c t i v i t y whereas the roots of plants exposed for 2 hours and held for 8 hours retained about 35-45% of the total  a c t i v i t y taken into the plants.  The transfer  of phosphate out of the roots was suppressed when the plants already contained adequate phosphate (5 ppm Experiment V).  S l i g h t l y more phos-  phate appears to have been transferred to the tops in older plants (48, 60 and 70 hour samples) as compared to the younger ones in each experiment.  Finally,  in each experiment the percentage of the total  count found in the stem tissue remained r e l a t i v e l y constant  (12-18%)  regardless of the holding time, age or P n u t r i t i o n of the plants; consequently large changes occurred in the root-leaf d i s t r i b u t i o n s which did not markedly affect  the percentages  in the stems.  Treatment of the plants with 50 ppm 2,4-D (Table IX) did not a l t e r the d i s t r i b u t i o n of phosphorus among the organs of the plant, and a l l of the same trends apparent in the controls were reproduced in the  42  Table stem  VIII. and  Percentage  root  tissue  of  distribution control  plants  of  the  total  harvested  at  activity  among  leaf,  various  times  after  32 initial  exposure  Expt. and Hrs, to Harvest  to P  Sampl i n g  % of  (hrs.)  5 ppm Expt.  11  Total  Activity  Times Leaf  Stem  Root  T/R*  2 14 60  17.5 19.8 46.6  14.0  68.5  1.46  11.7  68.5  1.46  11.5  41.9  2.39  0  18.9 21.3 25.0  16.8 12.3  64.3 66.4 58.1  1.55 1.51 1.72  17.4 17.2  35.8  2.79  34.1  2.93  5 ppm Expt.  Ill 16 60  5 ppm Expt. 10 5 ppm Expt. 10  IV 50 70  46.8  50  21.5 21.8  12.2  66.3  1.51  13.7  64.5  1.55  40.3  18.8  41.9  2.38  28.2  8.3 12.2 12.6  63.5 59.5 53.9  1.57 1.68 1.85  16.5 17.5 17.6  45.7 40.5 57.2  2.19  48.7  V'W 70  5 ppm Expt. 10  16.9  VI  5 0 ppm Expt.  I 3 20 48  28.3 33.5  50 ppm Expt.  I 1  10  10  37.8  10  20  42.0  4  70  25.2  *  Total  * *  Plants  activity/activity grown  in complete  in  roots.  nutrient.  2.47 1.75  Table stem  IX. and  Percentage  root  tissue  distribution of  treated  of  plants  the  total  harvested  activity at  among  various  leaf,  times  32 initial  Expt, to  exposure  and H r s ,  Harvest  5 ppm Expt.  to  P  ,  Sampling  % of  T o t a l A c t i V i t'y  T imes  11  4  (hrs.)  Leaf  Stem  Root  T/R*  2  18.7 35.0  13.5 14.2  67.8  1.47  6  50.8  14  20.2  30  32.6  10.9 14.8  1.97 1.45 1.90  60  21.7  14.7  68.9 52.6 63.6  2  22.0  14.8  63.2  1.58  16.3 17.4  61.4  1.63  66.2  1.51  76.9 53.3 73.0  1.30 1.88  1.57  5 ppm Expt.  111  4 8 3  16 30  22.3 16.4 10.2 26.0  60  9.0  12.9 20.7 18.0  50  41.6 43.2  20.0 20.0  38.4 36.8  2.72  70  21.2 20.0  12.7 16.0  66.1 64.0  1.56  1  40.0  19.6  40.4  2.47  3  27.2 26.1  8.6  64.2  8.0  31.3 36.0  10.9 15.5  65.9 57.8  1.55 1.52 1.74  48.5  2.05  1.37  5 ppm Expt, 10  IV 70  5 ppm Expt. 10  2.60  V**  50  1.51  5 ppm Expt. 10  VI  50 ppm Expt.  1  4  9 20 48  50 ppm Expt.  *  II  10  10  32.4  20.1  47.5  2.10  10  20  40.2  18.1  2.40  4  70  29.9  14.3  41.7 55.8  Total Plants  activity/activity grown  in complete  in  roots.  nutrient.  1.79  after  treated  plants.  Shortly  after  treatment  had  approximately  the  was  also  5 ppm  true  tribution or  60  while  higher  distribution hour  less  samples  activity  less.  5%  and  V),  70  hours  Cf  Percentage  and  acid  way  acid  soluble  trol  plants  Appendix  control  to  did  of  in which  the  and  insoluble  acid  various  The  and  and the  stem  only  roots  III.  in  change  the  total  the  roof-leaf  tissue  these  complete  not  controls.  large  but  the  plants  as  in  was  This dis-  equal  differences  occurred  In  leaves,  grown  total  in  the  and  11-25%  Experiment  IV  only  (5 ppm  distribution  to in  16  samples,  nutrient  activity  activity  is  times  The  given  was  fractions,  sampling  plant  all  II  the  the  in  Ex-  the  plants  between  acid  soluble  between  the  fractions.  whole  total  in  IX)  spraying.  basis.  the  toward  after  cpm/g  the  In  treatment  a  for  leaves  were  distribution  at  on  the  plants  Generally,  controls.  transported  insoluble  The  the  (Table  2,4-D  distribution  I.  percentage  Experiments  the  the  percentage  the  than  of  5 ppm  Experiment  between  was  When  periment 50  varied  slightly  the  for  same  with  in  is  partitioned  for  given  each in  Table  activity  in each  Table  expressed  X,  organ  of  of  XVII  these as  the of  conthe  fractions  a percentage  of  count.  experiments  increased plants  except  incorporation  aged.  The  the into  plants  in  50  ppm  acid the  Experiment insoluble  50  ppm  I,  the  compounds  Experiment  trend as  I were  was the observed  Table  X,  Percentage  soluble  and a c i d  various  times  distribution  insoluble  of  the total  fractions  of  control  exposure  to  P  activity  between  acid  plants  harvested  at  32  Expt. to  and  after  Hrs.  Harvest  initial  Sampling  %  11  Soluble  T/Sol.*  84.3 77.0  17.5 15.7 23.0  1.21 1.18 1.28  0 16 60  89.5 83.2 81.5  10.5 16.8 18.5  1.12 1.20 1.23  50 70  49.2  48.2  50.8 51.8  2.03 2.08  50 70  55.4 47.6  44.6 52.4  1.81 2.10  1  82.2  17.8  1.21  3 20  48  38.8 52.5 61.2  61.2 47.5 38.8  2.58 1.91 1.63  10 20  60.3 38.0  39.7 62.0  1.60 2.63  82.5  14  60 111  3 5 ppm Expt.  Activity  Insoluble  2  4 5 ppm Expt.  of Total  Times (hrs.)  5 ppm Expt.  .  IV  10  5  ppm Expt.  V**  10  5 ppm Expt.  VI  10 50 ppm Expt.  1  4 50 ppm Expt,  1 1  10  *  Total  * *  Plants  activity/activity grown  in complete  in soluble nutrient.  fraction.  46  Table  XI.  Percentage  soluble  and a c i d  various  times  d i s t r i b u t i o n of  insoluble fractions  the  total  of  treated  to  P  activity plants  between  harvested  acid at  32  Expt. to  and  after  Hrs.  Harvest  initial  exposure  Samp l i n g  .  %  of Total  Activity  T imes (hrs.)  Soluble  1nsoluble  T/Sol.*  5 ppm Expt.  11  k  5 ppm Expt.  11 1  2  81.4  18.6  6  81.6  18.4  }k  83.4  16.6  1.23 1.22 1.20  30  77.0  23.0  1.30  60  64.3  35.7  1.55  2  89.2  10.8  1.12  k  88.4  11.6  87.1 83.0 84.2  12.9 17.0 15.8  1.13 1.15 1.20  80.7  19.3  1.19 1.24  50.2  49.8 49.4  1.99 1.98  8  3  16 30 60 5 ppm Expt. 10  IV  5 ppm E x p t . V'~'" 10  5 ppm Expt. 10  50 70  50.6  50 70  48.5  35.7 51.5  1.55 2.06  1  78.0  22.0  1.28  3  31.1  68.9  9 20 48  47.5 74.1 75.6  52.5 25.9 24.4  3.21 2.10  10  56.8  20  59.8  43.2 40.2  64.3  VI  50 ppm Expt.  1  k  1.35 1.32  50 ppm Expt.  1 1  10  *  Total  * *  Plants  activity/activity grown  in complete  in s o l u b l e nutrient.  fraction.  1.76 1.67  hi  to  have more  the  other  fraction  to harvest  ration  growth  experiments,  insoluble time  rapid  into  Supplying  a n d showed  during  from  at the start  the plants  with  the experiment  decreasing  the course  3 t o 10 h o u r s  the insoluble  of  of  than  incorporation  the experiment.  considerably particularly  phosphate  prior  into  the  Increasing the  increased  fraction,  in any of  the  incorpo-  f o r the older  plants.  to treatment  (5 p p m E x p e r i m e n t 32  V),  slightly  suppressed  the  insoluble Similar  for  plants  the  older  than  crease  more  upon  10 h o u r s were  the total  into  Accordingly, as  the plants  count  insoluble the trend  activity  with  The e f f e c t  XI  compounds was toward  in opposition  plants. (l4-22%)  present  insoluble cases,  i n t h e same o r  50 h r . )  occurred  the holding  the results  Small  fraction  increases  occurred  However,  the presence found  time  these  of  after  less  obtained in the  up t o  increases  in the incorporation  a t 20 a n d 48 h o u r s  to that  into  in the controls.  50 p p m 2,4-D.  to untreated  XI  The l a r g e s t d e -  V, of  found  in the insoluble  decrease  resulted  (Experiment  and Table  into  and Table  l n most  compounds.  to that  P  f o r the c o n t r o l s ,  experiment.  insoluble  nutrient.  treated  by a marked  aged,  o f each  was p a r a l l e l  in comparison  followed  activity  were  the Appendix  more  of  taken.  As was t h e case  insoluble activity  the Appendix  of  the treatment  into  in complete  XIX o f  the plants of  the start  times  XVIII  incorporate  the controls  the d i s t r i b u t i o n  percent  to  incorporation  grown  Table when  with  at  in Table  5 p p m 2,4-D.  appeared  in a c i d  (9%)  plants  with  those  the incorporation  a t t h e two s a m p l i n g  are given  sprayed  compared  slightly  in  data  plants  compounds when  fraction  or d i d not alter  of  spraying.  insoluble  in the controls  activity of  50 p p m  48  Experiment  II  a n d a l l t h e 5 ppm e x p e r i m e n t s ,  in  the controls  D.  Percentage  stem  and root  When pressed are  as a percent  changed  the total  each  insoluble activity  with  of  fraction  count,  experiment  in each  the values  the percent  incorporation  the sampling  the percentage  experiments XIII)  into  more  percent The  the acid  among  leaf,  in the acid  time  organ  given  in Table  i n the stem  into  root  fraction  In of  XII  varied  and leaf  and experiment.  insoluble  is ex-  by  tissue every  the  leaf  increased with age.  (Table  is  than  I.  the insoluble  t h e amount  considerably  In  count  of  of  Within  however  experiment,  of  extent  tissue  the a c t i v i t y  3-4%,  tissue  50 ppm E x p e r i m e n t  distribution  obtained.  only  of  and to a g r e a t e r  leaf  centage  At  insoluble i n some  the total  tissue  Treatment porated  the  than  fraction  c a s e s when  insoluble  the c o n t r o l s ; the trend  activity  50 ppm ( T a b l e  than  XIII)  20 a n d 4 8 h o u r s  count  at  among  the plant  IV a n d V I ,  a slightly  in general  removed  more  M l ,  incorporated  marked of  II,  organs.  of  plants  greater  activity  contained  about  showed  the  This  increase  in Tables  as a  XII  and  or a smaller  no c o n s i s t e n t of  total  older  plants  XIll).  per-  trend. incor-  ones.  the decreases  found  associated with  The d e c r e a s e s  of  is expressed  t h e same  the leaves  the younger  were  activity  (brackets  and the stems whereby  tissue.  5 ppm 2 , 4 - D  with  percentage  the root  the root  sprayed  were  in the total  an a l t e r e d  mainly  due t o  insoluble  distribution less  incor-  49  Table and as  XII.  root  Distribution  of  the acid  tissue  of  control  a percentage  of  the total  Expt.  plants  insoluble  at various  activity  sampling  among  times,  leaf,  stem  expressed  activity.*  and  % of Total  Activity  Sampling Times  (hrs.)  Stem  Leaf  Root**  5 ppm 11  Expt.  2 14  60  3.2 2.7 7.1  1.6 1.4 0.9  12.7 (72.3) 11.6 (73.5) 15.0 (63.7)  1.8 3.3 3.8  1.8 2.2 4.6  6.9 (64.8) 11.3 (67.3) 10.3 (55.2)  30.8  33.0  6.3 6.1  13.7 (27.1) 12.7 (24.7)  8.2 12.6  3.3 4.8  33.1 (74.5) 35.0 (66.8)  6.4  3.2  8.2 (46.5)  11.3 11.5 12.8  3.8 7.8 5.3  46.1  12.7  5.0 7.3  22.0 24.3  5 ppm Expt.  I l l  0 16 60 5 ppm Expt.  IV  50  70  5 ppm Expt.  V***  50  70 5 ppm Expt.  VI  1 50 p p m Expt.  1  3 20 48 50 p p m Expt.  30.4 Sampling exper  correspond  in brackets  total  Plants  times  to  hours  after  spraying  plants  in each  iment.  Numbers the  v-Vo'c  20.7  II  10 20  * *  28.2  insoluble  grown  are the a c t i v i t y activity.  in complete  nutrient.  expressed  as a percentage  of  50  Table stem  XIII.  Distribution  and root  expressed  Expt.  tissue  of  of  the acid  treated  as a percentage  of  (hrs.)  plants  at  the total  activity  various  times  among after  spraying,  Total  Activity  Times Leaf  Stem  Root-  5 ppm II Expt. 2 6 14 30 60  2.8 9.7 3.0 7.1 3.6  1.7 3.1 1.2 3.0 l l . l  14.1 5.6 12.4  5 ppm Expt. 2 4 8 16 30 60  2. 2. 1. 1. 3. 1.  5 ppm Expt. 50 70  leaf,  activity.  % of  and  Sampling  insoluble  12.9 21.0  (76.0) (30.0) (74.5) (56.5) (59.3)  II 7.0 7.0 9.0 12.8 9.6 15.3  (64.5) (60.3) (69.5) (75.5) (. 6 1 . 1 ) (79.0)  16.8 14.7  (33.5) (30.1)  25.2  IV 27.2 27.2  5.8  6.7 10.0  3.8 5.6  35.9  (70.4) (69.8)  5 ppm E x p t . VI 1  7.6  3.9  10.5  (47.5)  50 ppm Expt. I 3 9 20 48  18.0 13.6 4.6 5.6  3.9 2.9 3.9 7.1  47.0  5 ppm Expt.  7.5  V*  50 70  36.0 17.4 11.7  51  Table  (cont.)  XIII.  .% o f  Total  Activity  Leaf  Stem  Root*  13.9 13.1  6.1  23.2 20.7  5.0 ppm Expt. II 10 20  *  Numbers the  Plants  little  brackets insoluble  grown  poration  E.  in  total  into  change  in  the in  and  ethanol  In  the  5 ppm  to  of  the  compared  to  controls.  stem  the  as  acid  insoluble  insoluble  Experiment  IV,  amounts  components  in  the  organs.  root  85-90% o f  samples  insoluble  three  the  was  not  experiment,  the  treatment  activity  the  root  noted  in  above  in  the  root  activity  of  acid  ethanol  The  and  leaf  between  acid In  of  tissue,  the  with  ethanol  (Table  insoluble  soluble  fraction and  stem,  insoluble  and  the  ranged  the  ethanol  XX,  Appendix).  fraction  appear  found insoluble  fraction in  percentage from  was  ethanol  insoluble activity  the  increased  insoluble  ethanol  consistent  tissue  acid  the  total  respectively.  activity  a percentage  fractions  different  and  as  fractions  of  95% a n d  expressed  insoluble  consist  about  activity  nutrient.  of  soluble  the  activity.  complete  the  Distribution  are  composed  the of  35-80%.  leaf  ethanol In  this  insoluble The to  be  changes mainly  derived  from  F.  its  ethanol  Percentage  stem  and  the  count.  of  In  acid  leaf  tissue  the  gives the  all  less  cases,  the  changed  in  -P  roots  by  to  plants,  except  the  acid  soluble  as  only  the  plants  only  with  8-10%  prior  to  increase  as  the  activity  among  each  compared  a  soluble percent  leaf,  of  the  to  total  roots  percentage  among  retained in  the  another,  but  experiment.  plants  insoluble  This  activity  the  time  increased  to  The  I,  The  sampling  phosphate  20-22%.  as  aged.  one  within  acid  Experiment  plants  exposure. by  ppm  from  slightly  of  expressed 50  considerably  the  nutrient  previously  the  distribution  activity  varied  Supplying in  the  control  soluble  stems  of  tissue  XIV  organs  components.  distribution  root  Table  insoluble  the  soluble  (Experiment activity  difference  is  activity  IV)  was  grown  seen  undoubtedly  re-  32 lated from  to the  located of  the  soluble upward  dilution,  uble  fact  the  P  fraction.  and  so  activity  that  it  that  but  lost On  the  remained in  the  22-25%  the  other  mainly  tops  less  to  hand,  in  these  nutrient  the  was  derived  activity  was  trans-  soluble  plants  insoluble  less  solution  fraction  contained  activity  than  4-8%  those  because less  grown  solin  -P  nutrient. Treatment distribution roots 50  ppm  with  as  contained treatment  5 ppm  compared more  or  2,4-D to  the  (Table  the  controls,  leaves  accumulation  of  XV)  less  acid  of  in  general  although the  soluble  total  in  did some  not  alter  cases  activity.  compounds  occurred  the  the With in  the the  53  r o o t s and l e a v e s a t 20 and 48 hours a f t e r s p r a y i n g .  The stems c o n t a i n e d  about the same o r a s l i g h t l y h i g h e r percentage throughout ments.  the e x p e r i -  54  Table and as  XIV.  root  Distribution  tissue  of  control  a percentage  of  the  Expt.  of  the a c i d  plants  total  various  activity sampling  among times,  leaf,  stem  expressed  activity.*  and  Sampl i n g  at  soluble  % of  Total  Act ivi ty  Times  (hrs.)  Leaf  Stem  Root  14.3 17.0  12.3  14  10.3  55.9 57.0  60  39.6  10.7  26.7  17.1 18.0  15.0 10.0  57.4  21.3  12.2  55.2 48.0  16.0  11.1 1 1.0  22.1 21.4  14.9 9.2  8.6 8.9  3 K 9 29.5  33.8  15.6  32.8  3 20  17.0 16.8  4.4  17.4  4.6  48  20.6  7.2  31.1 33.4  11.5 10.1  23.7 16.2  5 ppm Expt.  1I  2  5 ppm Expt. 0  111  16 60 5 ppm Expt.  IV  50 70  15.8  5 ppm Expt.  V**  50 70 5 ppm Expt.  VI  1 5 0 ppm Expt.  1  5 0 ppm Expt. 10  11 25.1 11.7  20 *  Samp]ing  t imes c o r r e s p o n d  to hours  e x p e r imen t . * *  Plants  grown  in complete  nutrient.  after  spraying  plants  in  each  Table and  XV.  root  expressed  Expt.  Distribution tissue  of  of  treated  as a percentage  the acid plants  of  the  and  at  soluble  activity  various  times  total  among  after  leaf,  stem  spraying,  activity.  % of  Total  Activity  Sampl i n g T i m e s (  h r s  -)  5 ppm Expt.  Leaf  Stem  Root  53.8 45.0  II 15.8  11.8  25.4  11.2  14  17.4  30  25.6 18.0  9.7 11.8  2 6  60 5 ppm Expt.  3.7  56.3 39.6 42.6  Ill  2 4 8  20.0  13.2  19.5 14.6  14.5  16  30  8.4 23.0  60  56.0 54.4 57.2 64.1 43.6  7.2  15.3 10.5 17.6 15.8  14.2 16.1  14.1 12.4  21.9 22.1  14.6 10.0  8.8 10.4  41.0  32.5  15.7  29.8  3 9 20  9.1 12.7 26.6  4.6  17.4 29.6  48  30.3  8.5  5 ppm Expt.  IV  50 70 5 ppm Expt.  57.7  V*  50 70 5 ppm Expt. VI 1  28.1  50 ppm Expt.  1 5.2 7.0  40.5 36.8  Table  XV.  (cont.)  % of  Total  Activity  Leaf  Stem  18.5  14.1  24.2  27.4  11.6  20.8  Root  50 ppm Expt.  II  10 20  "  Plants  G.  grown  in  Distribution  complete  of  activity  Standardization described  under  isolation  of  within  the  of  the  picric  Methods  and  Materials.  any  individual  phage,  when  many  it  give  reproducible  did  nutrient.  phosphate  acid  compound, esters  acid  solvent This  of  the  fraction  system  system  except  occurred  separation  soluble  did  possibly  in  the  was not  as  allow  inorganic  same  standard  done  the  phos-  sample.  However  compounds  into  the 32  four was  regions studied  plants  Leaf  given  in  comparing  mainly  by  Table the  XVI.  The  scans of  reference  to  pattern  extracts  these  of  from  incorporation treated  and  of  P  untreated  regions.  tissue-  Figures fractions  of  3a,  4a,  and  the  leaves  5a of  show  typical  controls  from  scans  from  the  acid  three  different  soluble  experiments. 32  It  was  apparent  incorporated  that  into  there  was  a wide  soluble organic  variation  compounds  from  in  the  one  amount  sampling  of  P  time  or  Table  XVI.  developed  Location in  the  of  picric  standard acid  or  phosphorus isobutyric  compounds acid  R  solvents.  *  Picric  at  UTP  1  1sobutyr i c  .13  origin  ATP  .008  .50  UDP  .04  .31  A DP  .06  .96  Phytate  .09  F-6-P  .27  GMP  .33  G-l-P  .39  G-6-P  .41  1 1  chromatograms  Sol v e n t  Compound  Reg i o n  P  on  at  origin  .42 FDP  .42  R-5-P  .47  P-Chol  11 1  .71  ine  .83  PGA  IV *  PV  Rp  equals  by  PO^""3.  butyric);  the  1 .00  3  distance  travelled  Reproducibility:  region  II  ±.03;  by  region  region  III  the I  compound  £.03 ±.05.  /distance  (picric),  i.05  travelled (iso-  58  experiment traction a  to  and  similar  at  These  concentration  (regions  which  sufficient lation  This  distribution  compounds rate  another.  the  1-1  changed  from  differences  completely  I I).  were  taken  sampling  to  hydrolysis  during  the  since  duplicates  always  showed  effect  trends  to  the  the  on  the  While  appeared  time  in  whole  sufficiently  holding  ex-  organic  aged.  whether it  and  observed  they  another, were  IV)  were  as  determine  times  of  (region  activity  time  sampling  any  to  consistent  incorporated  between  overshadow  due  orthophosphate  No  not  one  not  procedures  between  plants  samples  was  popu-  that  way.  large  to  the  amount  of  exposure  showed  only  i ncorporat ion.  Plants inorganic result lower  about  same  under  other  (Figure  than  the  3c)  the  due  to  from  of  This  dilution  plants  in  at  to  fraction of  the  was  between  sprayed the  Figure  incorporation  phosphate  compounds,  similar  difference  previous  soluble  c o n c l u s i o n was  inorganic  there  difference  acid  presented  phosphorus A  the  harvested  amount  controls.  background.  The  are  3a,  the  and  nutrient  of  the  isotope  leaves.  rather  This  than  a  rate.  taken  exposure,  the  the  in  undoubtedly  samples  Figure  areas  complete  phosphorylation  before  in  in  phosphate  was  Two  in  grown  these  between  time  as  and  into  c.  by  with  indicates  The  incorporation  samples duplicate  from  l - l l l  comparing  small that  under  hour given  sample as  was  the  in  the  the  than  other in  the  plants  samples  and  shows found  relative peaks  fluctuations  treated  control  control  first  those  one  2,4-D  the  regions  reached  peaks  more  two the  3b  5 ppm  disregarding  comparison slightly  same  with  due  of to  sample controls. was  greater  Indicates  Figure  4b.  Acid  soluble  fraction  of  leaf  tissue  from  plants  sprayed  with  5 ppm  2,4-D.  ON VjO  66  that  in  this  acid  soluble,  same  result  spraying,  as  as  pattern  with  in  Figure  which  showed  Stem  tissuecan  more  be  in  an  1-1  grown  I I and  ppm  2,4-D  stem  but in  treated  the  and  7b  5 ppm  7a.  In  6a  pattern  slightly  and  from  7a  found,  The  hours  after  However the  in  treated  pattern  did  not  as  treated  of  as  most and  incorpo-  the  were  and  larger sampling quite some  from  the  in  to  of  inorganic  another The  be  within  stems into  of  regions  respect.  involving  controls  treatment to  of  although  experiments the  controls.  pattern  to  this  ex-  found  to  proportion time  or  for  were  leaves,  similar.  two  correspond  fact,  shown  incorporation  roots  the  rate  compared  of  a  is  the  samples  that  one  in  alter  that  experiments,  was  general  8a  samples and  between  and  showed  leaves  14  and 4 b .  compounds  contained  samples  2,4-D  4a  compounds,  resembled  the  both  this  6a,  nutrient  treatment.  into  system.  organic  represent  with  or  generally  occurred  from  rate  organic  former  incorporation  following  Figure  acid  Figures  duplicate  and  Figures  in  P  5 ppm e x p e r i m e n t ,  Occasionally,  samples  complete  6b  into  b.  from  differened  Figures plants  50  Variations  experiment  plants  picric  of  differences  the  the  activity  characteristically phosphate.  distinct to  rate  compounds  comparing  by  seen  the  by  respect  and  5a  increased  in another  no  incorporation  ample  activity  seen  with  an  phosphorus  were  revealed  of  It  be  there  Spraying  was  obtained  can  plants  ration  there  organic  was  experiments control  case  the  only case  given  changed in  in the  general.  OO  Figure  8b.  Acid  soluble  fraction  of  stem  tissue  from  plants  sprayed  with  50 ppm 2,4-D.  73  The  same  situation  was  Figure  8a  and  lation  or  i n h i b i t i o n of  pattern  Root  of  Figure  found  holding scans  roots  that  time  from  served  found  in  the  stem  in  found  in in  acid  tissue  sampling other  in  level,  gave or  no  of  for  example  indication  any  marked  compare  of  stimu-  change  in  the  tissue.  of  of  of  the  amount  9a  of  treatment  proportion  Figures  9a,  control to  10a  Plants in  could  and  but  their  appropriate  and  b and  Figure  samples  inorganic with  the  as  from  a  10a  phosphate higher  was  diminished show  as  the  representative was  was  generally  less  than  nutrient  con-  in  complete (Figure  the  and of  b.  as  The  either  sprayed  plants  can  be  ppm.  50  seen  large  treatment  considerably  concentration  for  only  with  ob-  Appendix).  17.  scans  controls,  result  plants  radioactive  variation  between  and  it  11a  fraction  found  of  Some it  grown  this  be  but  plants.  another  incorporation root  large  fraction,  organs.  2,4-D  a  soluble  phosphate  Figure rate  of  time  differences  5 ppm  some  result  increased.  the  the  relative  their  inorganic  with  example  change  in  root  marked  treated  contained  one  only  No  a  treatments  commonly  was  the  from  tained  as  ppm  50  phosphorylation  incorporation  orthophosphate  the  The  8b.  the  tissue-  The  for  at  was Here  increased  (Figure  11a  and  Nucleot ides-  Separation  in  the  isobutyric  acid  system  did  not  prove  to  be  too  b).  Figure control  10a.  Actigraph  scan  of  picric acid  chromatogram;  acid  soluble  fraction  of  root  tissue  plants.*  origin Discontinuity  indicates  that  the  vertical  scale  is  doubled.  from  Figure  10b.  Acid  soluble  fraction  of  root  tissue  from  plants  sprayed  with  5 ppm  2,4-D.  *  Discontinuity  indicates  that  the  vertical  scale  is  doubled. vo  80  informative. the 5  scans  ppm.  UTP  Only  from  One  or  H.  were  treated  Distribution  The  stems  hours  were  leaves.  The  8  and  of  cut  areas  leaves  of  and  was  found  not  of  the  exudate  was  plants  which base  immediately  spotted  using  the  picric  solvent.  one  given  active with  in  Figure  component  5 ppm  2,4-D.  of  12. the  and in  any  were  had  on  at  be  All  and  of  the  plants to  ATP  in  all  treated and  samples.  activity  in  the  with  possibly  No  marked  to  areas  compared.  exudates  been the  exposed point  it  the  was  of  scans  for  2 hours  released which  of  from were  obtained  untreated  and  attachment  orthophosphate  from  distinguished  other  chromatograms  Inorganic exudates  the  relative  c o l l e c t e d as  and  could  controls  xylem  or  faces  acid  the  in  of  ADP  tagged  activity  plants at  to  between  untreated  activity  roots  corresponded  Phytate  differences when  the  area  UDP.  two  was  plants  the  and  the  for  primary  cut  sur-  developed  were the  held  like  only  the radio-  plants  treated  Figure  12.  Actigraph  scan  of  picric  acid  chromatogram;  xylem  exudate-  82  DISCUSSION  A.  Phosphate  From control of  uptake  Table  plants  III in  50-60 h o u r s ,  for  the  during  zation  of  The  while  2,4-D.  uptake  Within  a plant  not  of  the  first  rather  than  over  a  2,4-D  dry  and  period  may  VII.  be  a  moisture  hours  doubled a  uptake  over  period  increase  r e f l e c t i o n of  has  responded  of  content  day  old  to  after  could  Also,  not  the  in  the  a  the  by  period  of  60  hours  rate  gradual  of u t i l i -  relationship  spraying, It  lead  to  is a  P  uptake  still  between  level can  possible  cumulative  of  be that  effect  on  P  days.  c l o s u r e would reducing foliar of  (86).  bean The  treatment  tend  to  plants use of the  much  continued  diminish  transpiration  a p p l i c a t i o n of  confounded  plants  direct  growth-stimulatory  unchanged.  stomatal  treatment  weight,  14  The  i n d i c a t e any  i n c r e a s e s by  (l04).  after  nutrient  auxin  on  early  not  remain  the  the  hours  nutrient;  in Table  50-70  by  uptake  alter  21  -P  the  ion  within  complete  of  phosphate  phosphate.  inhibited or  by  total  increase occurred over  a p p l i c a t i o n of  uptake  inorganic  such  and  stimulation  magnitude  no  the  approximately  do  the  effect  that  results  growth  An  seen  experiments  reported  available  stimulated,  be  some e x p e r i m e n t s  present  phosphate  can  three  experiment  uptake  does  it  by  to  same  grow  at  than  weight  results. the  thus  2,4-D  more  fresh  and  as  well  the  100  ppm  1-2% therefore,  Plants  grown  in  those  grown  in  for  1-2  weeks  83  in  the  -P  in  fact  solution.  appears  Further to  the  level  growth, from  did  nism  of  of  followed 2,4-D and by  was  the  Both  inhibit seedlings  gave  treatments  by  Cooke  metabolism  As  yet,  ppm  this  P  as  uptake  50  In  is  a  any  therefore  mentioned  was  not  respect has  undoubtedly than  acute  while  bean  been  and  above.  directly  treatment,  uptake  similar  same  rate  These  of  no  the  Salisbury  leaves.  the  not  related  inhibiting  seedlings  lowered  related  to  a  difference  in  the  differ  by  a  50  greater  ppm  sensi-  basic  mecha-  effect.  the  and  that  was  uptake  phosphorus  rather  or  Ross  more  in which  sunflowers  in  to  difference  treatments  (4)  deficiency  uptake.  less  Dyke  bean  2,4-D  2,4-D  by  lead  of  This  the  have  that  not  (10).  tivity  to  evidence  sunflower  spray  Phosphorus  in  changes  adequate  in  as  the  the  a  initial  explanation  controls,  of  to  intact  general reactions  has  been  Le0  stimulation  that  uptake  hydrolyzable  reflect  from  pattern,  mineral  elongation  (101)  resulting  uptake  (34) of  response  of  Avena  found beans,  transient  for  Ball  levels  and  of  stimulation  2,4-D  presented  by  coleoptiles,  phosphate  of  after  within such  a  the  plant.  response  pattern.  The increases  fact in  that the  after  spraying  ation  between  cannot  reach  lation  must  foliar  rate  is of the  the  of  P  uptake  interest  leaf roots  originate  applications  and in in  in  root so  the  by  of  roots  itself. since  short  a  leaves,  the time  the  auxin  within This  possibly  as  l i t t l e  indicates  2,4-D (35,  caused  molecules 50, in  68).  the  a  transient as  one  close  hour  associ-  apparently Such  same way  a as  stimuchanges  84  which bean  cause diurnal roots  (49),  synthesis  B.  of  rate  in  the  connection  translocation  with  transient  of  phosphate  increases  in  across photo-  translocation  data  movement  the  and  in  (120).  Phosphate  The  variations  presented  of  of  in Table  phosphate  net  upward  out  of  VII  can  the  be  roots  translocation  of  used  into  to  the  inorganic  evaluate stem,  the  which  phosphate.  rate represents  If  T  =  32 the  total  tained has  amount  by  the  P  roots  occurred,  This  of  then  equation  taken  at R =  end  T  cst»R»r«t  -  rearranges  the  T/R  values  nutrient  are  plotted  the  youngest  (Figure it.  13),  Since  while  the  same A  shows  among  pattern,  similar that  the  treatment  distribution  all  as  time  t,  and  during  where  r =  the  R =  the  activity  which rate  re-  translocation of  translocation.  a  5 0 ppm  be the  seen  along  previously  harvest, which  not  values Table  sprayed the  ones  grown  samples) alter  for  the  these  in  -P  representing  a y-intercept  hour  did  T/R  the  has  (30-70  from  plants  lie  to  2,4-D  the  (1)  controls  line  plants  organs,  cst-r.t  for  hours  older  values  1 +  VIII)  along  can for  =  the  with  the  plot the  vs. lie  a  roots,  to:  (Table  plants  of  the  the  T/R When  into  all  of  1  lie  above  percentage plants  follow  IX. with  line,  5 ppm 2 , 4 - D  the  effect  of  upward  of  (Figure aging  14)  being  removed.  The  data  indicate  therefore  that  the  rate  translocation  Figure  13.  T/R  vs.  hours  to  harvest  for  control  plants  of  two  age  groups.*  1  0  1 2  1  k  Hours  to  •  Younger  X  Older  1  »  :—•  6  8  10  Harvest  * T/R = t o t a l a c t i v i t y / a c t i v i t y in roots. Younger s a m p l e s ; \ o l d e r g r o u p i n c l u d e s 30-70 h o u r s a m p l e s .  group  includes  1-20  hour  86  'Figure  14.  T/R  sp,rayedcwitHi.5  *  T/R  =  'samples;  total older  vs.  ppm  hours  to  harvest  for  plants  of  two  age  groups  2,4-D.*  activity/activity group  includes  in  30-70  roots. hour  Younger samples.  group  includes  1-20  hour  87  of  phosphate  each  experiment.  available this  trend  or  low  added  to  as  132%  the  the  activity  probably  plants.  younger  older  and  enhance  the  Accordingly,  spray  solution  activity  as  the  The  when  used  this  the  controls,  to  2,4-D  5 ppm  the  a  period  decrease  either  did  treatment  in  effects  450 the  ppm  from  tops.  The  the  Fe  5 ppm 3923  additional  and  over  of  in not  affect  removed  the  plants.  increased  but in  ppm  stimulatory  84).  plants  aged  related  Fifty it.  controls;  of  plants  accentuated  activity/3  of  is  the  of  2,4-D  as  Fe  leaves  EDTA  was  Experiment  I,  the  (Table  to  4159,  increase  roots  applied  now  and  II) was  associated  contained  stems,  140%  125%  and  respectively.  87%  Both  the  hypothesis 2,4-D  interaction  that  promotes  tion to  5 ppm  a decrease  much  to  (83,  total  to  with  the  in  the  known  levels  average ie.  is  change  slightly  between  Iron  increased as  This  phosphate  difference  at  normally  of  any  growth more  entering inhibitory  increased  with  age  and  the  stimulation  due  to  active  participation  phosphorus. effect  metabolic  on  This  in  foliar by  response  translocation  activity  effect  the  roots  the is  per  of  iron  application roots  not se.  and/or  support  in  of  the  may  5 ppm  distribu-  considered It  the  to  be  due  represent  decreased  activity  in  32 the  leaves.  Fang  and  beans. upward anism  Similar  Butts  (4l)  changes following  Concentrations translocation is  trations.  undoubtedly  in  of  of not  P  the  distribution  10-100  2,4-D in  the  ug  which  barley same  as  that  P  applications  retard  (110)  of  and  P  uptake beans  which  were  found  of  2,4-D  can  also  (98),  operates  but at  by  to retard the  low  mech-  concen-  88  As phorus  would prior  to  plants.  This  location  is  lower pool  rate of  be  expected exposure  finding  related was  not  inorganic  Maizel  role  in  amount  and  a  by  has  the  in  been  it  organic  used  compound  the  solvent  showed  concluded ward  C.  that  either  pound (s)  only  in  not  taken  during  which  poration.  When  is  made,  not  in  the  all  cases  been  the  xylem  suggested  Loughman  and  compound  similar  system.  Rp  in to  room  the  However,  T/Sol.  plant;  of  plants  (Figure  accompanied  temperature  incorporation plot  or  insoluble  Sol.  =  in  the  the  large  that  it  of  may  is  a  play  organic  xylem and  translocated  acid  small The  unidentified  labile  some  a  exudates.  highly  picric  barley  found  (70)  radioactive  follow  this both  where,  soluble  occurred; values  In  the  investigation,  by  1 -|- e s t - r - t  T/Sol.  15).  of  The  exuit  is  up-  com-  solvent.  compounds  =  has  trans-  plant.  exudates  this  phosphorus  at  low-P  upward  the  barley  the  alone  than  of  Russell  of  into  older  in  beans form  rate  phos-  tissues.  a  bulk  rates  because  present  becomes  a  treatment  the  the  adequate  within  in  into  (l)  the  differences  has  had  that  orthophosphate  this  Incorporation  activity  same  stable  Equation  t,  in  fact  phosphorus  standard  contained  translocation  identified  and  (74)  which  5 ppm  the  phosphochol ine  dates  the  translocation.  single  using  lower  concentration  phosphate  phosphorus of  showed  altered  Benson  plants  emphasizes  to  Phosphochol ine by  (4-9),  and  (Table  equation, treated  activity  r = X)  the  vs.  but and  T =  the after  rate  hours  the  of to  younger  untreated  total a  time  incorharvest ones  plants  do  Figure  T/Sol.  15.  vs.  hours  to  harvest  for  control  plants  of  two  age  groups.*  J  2.0 T/Sol.  1.0  Hours  to  Harvest  \  ^ ( S ' 7 T  S  1  .Deludes  ^ i  1-20  3  '  y / a c t i v i t y in s o l u b l e f r a c t i o n . Younger group s a m p l e s ; o l d e r group i n c l u d e s 30-70 hour samples.  a c t i v i t  hour  Figure 5  ppm  16.  T/Sol.  2,4-D.*  vs.  hours  to  harvest  for  older  plants  sprayed  with  91  (Table  XI  soluble of  the  the  and  Figure  compounds insoluble  at  16),  the  older  the  same  rate.  compounds  variation  in  the  exposure  to  harvest  from tween  the  roots  necessarily  and  younger  in  a  as  larger  to  of  between  P  while  a more  result  a balance  variable  soluble  Also,  led  a  incorporated  A more  group.  always  leaves  result  and  ones  activity rate  pool  of  balanced  soluble  account the  for  period  distribution  translocation,  the  in-  turnover  might  increasing  into  it  and  did  be-  not  insoluble  frac-  t ions.  The  change  observed  by  Cherry  et  aj_.  (23)  in  related  to  the  fact  that  found in  part  nutrient.  These  in  authors  the  plants  with  peanut the  h a v e made  age  seems  cotyledons, plants  the  were  pertinent  to  parallel  although grown  here  in  that  it  is  phosphate-free  statement  that:  . . . t h e phosphorus pool w i t h i n the c e l l s of peanut cotyledons r a p i d l y d i m i n i s h e s d u r i n g g e r m i n a t i o n and growth o f the plant. T h i s l e a d s t o a much e n h a n c e d r a t e o f phosphorus-P i n c o r p o r a t i o n i n t o RNA w i t h a g e . These r e s u l t s p o i n t up the d i f f i c u l t y of attempting to judge rates of synthesis of material in d i f f e r e n t t i s s u e s without knowing pool sizes or the e x t e n t of d i l u t i o n of the a d m i n i s t e r e d isotope. The d i s t i n c t i o n between a s m a l l a c t i v e pool and a l a r g e n o n a c t i v e pool of the d i f f e r e n t m e t a b o l i t e s in the p l a n t cell (12) further complicates this issue.  Both sponses  the  which  85% o f  this  terial  and  IAA,  inhibitory  has  amount  of  stimulatory  appear  to  be  fraction  of  the  therefore  is  considered  depending  thesis  and  been  on  the  shown  stimulation  related root  to  concentration, (59, by  103). 5 ppm  the  tissue to  The 2,4-D  levels acid  was  of  2,4-D  insoluble  also  ethanol  be  primarily  RNA.  can  stimulate  or  present in  the  data intact  produce  re-  fraction.  About  insoluble  ma-  The inhibit  fact  that  RNA  syn-  indicate  that  plant  small,  is  the and  92  primarily not  related  observed  increase the  in  an  in every the  apparent  synthesis  to  size  effect  case of  but  the  to  the  does  On  occur  roots.  appear  soluble  incorporation.  appears  on  More  as  a  phosphorus  the  other  in  both  the  be  inhibited  trend pool  hand,  leaves  incorporation in  may  data.  have  marked  and  the  was An  suppressed  i n h i b i t i o n of  roots  at  the  50  RNA  ppm  1evel.  Aboveground time the  as  roots  tops  (48,  growth are  D.  136).  retardation  emphasizing  the  The  of  standards  the  (19, the  70,  scans are  especially pecially IV)  tides  present  concentrations  2,4-D  of in  of  can  the be  support  change or  IAA  2,4-D  of  at  the  a part  same  is a p p l i e d in  treatments  data of  to  overall  foliar  present  as  at  result  which  the  stimulation  intact  The  and  acid  by  the  restimu-  plants.  main to  I  into  be:  I)  and  by  results  pentose  ribose;  the  of  is  III)  the  acid  by  the  this  regions  and  basis system  marked  phosphates,  phosphates,  from  the  on  triphosphates,  hexose  triose  on  with  four  d i -  and  revealed  parts  obtained  separated  isobutyric  as  major  nuceloside  II)  ADP  fraction  four  components  uridine;  fructose  region  soluble  previous  orthophosphate. in  root  divided  and  considered  adenine  inorganic  same  used  135).  glucose,  2,4-D  the  results  no  compounds  system  100,  when  applications  These  composition  acid  show  root  at  of  growth  or  Also  importance  soluble  picric  increased  (48).  mechanism  Acid  can  undergo  stimulatory  latory  parts  the  system.  other  es-  and nucleo-  93  The the  distribution  controls  within While  an a  occurred  in  be  or  and  have  phosphate external large  (12) in  in  potato  in  within all  out, and  the  the  changes  of  established tissue.  phosphate  first  nonmetabolic  pool.  is  be  Depending  a  of  the  fraction  sampling number  were  concen-  plants  two  rates  passing  also  of  Bieleski  pools  metabolic  and  harvested  pools.  relative  sources.  large  in  of  times  of  and  variations  slowly  on  have  phosphate  small  before  all  treated  presence a  at  extraction  from  soluble  One  enters  the  could  the  or  soluble  encountered  taken  sources  acid  could  during  samples  size  the  organs  variations  hydrolysis  Possible  phosphorylation Laties  of  ruled  time.  same  These  duplicates  same  activity  the  amount  cannot  the  not  experiment.  small  tration  at  was  of  of  inorganic  pool  into  into  the  which other  importance  of  pool  32 sizes not  on  the  closely  rates  of  reflect  ward  translocation  sent  the  (ll)  have  movement found  incorporation,  synthetic would  of  that  7  rates.  also  only  add  after  distribution  Different  to  a portion  hours  the  these of  the  rates  of  P  may  of  upward  variations  since  fraction.  Biddulph  exposure  of  beans  having  or or  they  downrepre-  et  3  may  a 1.  trifoliate  32 leaves  to  dicating first  P  there  that  a  entering The  soluble  was  portion the  absence  of  of  tracer  any  consistent  following  above  factors.  Whenever  5 ppm  level  can  stimulate  While  it  difficult  is  the  activity  in  moved  the  to  the  primary stem  leaves,  apexes  in-  after  leaves.  of  fractions  a peak  to  changes  treatment  differences  could were  incorporation compare  the  in  the  be  found  and  results  the  composition  partly  related  they  indicated  higher  obtained  level with  of  the  to  the  that  inhibit different  the it.  plants to  and  those  different  reported  by  treatments,  these  Berezovskii  and  Maciejewska-Potapczyk trend or  toward  toward  beans  as  cannot the  found  roots  used  case  it  rather  Within  the  no  1 ittle  grown  to  where  than  soluble  5 ppm  in  the  fraction,  stems.  leaves  (10)  by  and  do  not  Ormrod  and  appear  more  Humphries show  and in  similar  the  and clear  Williams  the  stems  (92) of  (4l).  complete  2,4-D  reduced  was  the  effects  results  found  synthetic  represented  incorporation  in  Butts  under  indicate  a  the  incorporation  Fang and  probably  tissue  that  by  Kurochkina  However  incorporation  plants  be  one  increased  less  With  (54).  treatment  nutrition rates  increased a  greater  rate the  of  the  soluble  synthesis  in  the  of  present  soluble  retention  dilution  recorded  of  the  of  the  roots  of  compounds. activity the  isotope leaves  in  ion  insoluble  or  procedure  by  In the the  compounds.  was and  so  great  very  95  SUMMARY  A tions  study of  plant,  mechanism  thus  the  gain  whereby  Bean  of  carried  on  2,4-D  and  sprayed  was  plants  with  treated  and  to  uptake  determine  and  information affects  2,4-D  grown  5 or  out  in  ppm  50  untreated  which the  could  growth  when  plants  effect  distribution  phosphate-free 2,4-D  the  were  be  of  or  they  of  of  low  concentra-  phosphorus used  to  intact  by  the  describe  bean the  plants.  complete  nutrient  were  14  days  old.  immersed  at  various  were  The  times  roots up  to  32 70  hours  for  1-2  grown.  after  spraying  in a c o m p l e t e  hours,  and  then  returned  At  end  of  holding  the  a  separated  into  roots,  activity,  acid  insoluble  within  the  recorded  soluble  for  each  Control  plants  showed  another  ing  of  available  soluble  complicated tively The l)  small  treatment No  which  lasting  and  the  revealed  these  effects effects effect  variations were  by  pools.  and  nutrient plants  and  in  were  leaves.  distribution paper  on  In  phosphorus  can  be  the  containing which  harvested Acid of  P  they  were  and  soluble activity  chromatography  of  sampling  related  changes  uptake, as  to  in  general  summarized rate  one  Interpretation  variations. on  from  probably  phosphorus  phosphorus by  the  petioles,  activity, as  same  solution  were  organ.  to  acid  the  period,  plus  fraction  periment pool  stems  to  nutrient  a  the of  the  time  and  gradually relative  treatment treatments  translocation  or  ex-  diminish-  sizes effects had  of was  rela-  incorporation.  follows:  phosphate  uptake  was  observed  for  either  treatment.  tained  and were  2)  The r a t e  the  age o f  of  Initial followed  upward  the p l a n t ,  by l e s s  3)  a s compared  the  plant  Fifty  incorporation  was unchanged  ppm r e d u c e d  48 h o u r s 4) A t  after  and leaves  5)  The c o m p o s i t i o n  6)  While  the total  the lower  after  in t h i s  toward  more  and less  had reduced  of  the acid  by  with  had no  organs. insoluble  the  fraction  of  growth-regu1ator.  fraction  b y 14-22%,  incorporation  into  in the case of  incorporation  at either  soluble  20 a n d  at  acid  in-  the leaf  tissue  t h e 50 ppm l e v e l  bending  and leaf  of  fractions  showed  no c o n s i s t e n t  concentration.  5 ppm h a d n o v i s i b l e e f f e c t  elongation  treatment  concentration  the  of  affected  spraying.  controls  characteristic  and less  acid  level  activity  in the roots,  20 h o u r s  from  into  among  not main-  as the c o n t r o l s .  after  The h i g h e r  activity  was a t r e n d  by  changes  was reduced  were  treatment.  compounds  the roots  at  the total  5 ppm t h e r e  soluble Both  of  of  plants  uptake  to controls,  Fe EDTA.  on the d i s t r i b u t i o n  The r a t e  o r t h e same  translocation  5 ppm 2,4-D o r 5 ppm p l u s effect  i n c r e a s e s by t r e a t e d  on growth,  the petioles  a n d some  the higher  level  retardation  of  caused  stem  expansion.  32 The  results  indicate  tribution  between  by  application of  foliar  latory to and  the leaves  and i n h i b i t o r y  be r e l a t e d leaves.  a pattern  and roots  5 ppm 2 , 4 - D ,  levels  of  to the synthesis The major  whereby  part  of  the balance  i s upset  in f a v o r  in P of  b u t n o t b y 50 p p m .  the auxin of acid the acid  produce  insoluble insoluble  activity  the  Both  responses compounds  disroots stimu-  which  appea  in the  roots  of  these  97  tissues  were  also ethanol  Actigraph corporation  of  phosphates)  in  i n s o l u b l e and  scans of  picric  activity  into  the  and  stem  tissues  plants  from  the  leaf  acid  is  believed  chromatograms  organic  compounds than  to  be  RNA.  i n d i c a t e d more  (nucleotides in  the  roots  and up  insugar  to  hours  8  32 after  removal  Plants  of  grown  the  in  complete  P  nutrient  solution.  prior  to  treatment  and  exposure  32 to  P  grown  ,  took  up,  translocated,  in phosphate-free  organic  compounds  The  only  was  and  nutrient.  retained Also,  less  phosphate  incorporation  than  into  those  soluble  suppressed.  radioactive  compound  in  xylem  exudates  from  treated  and  32 untreated phosphate.  plants  10  hours  after  their  initial  exposure  to  P  was  ortho-  BlBLIOGRAPHY  A l e x a n d e r , A l e x G. 1965. a c t i v i t y of immature of indole-3-acetic ac maleic hydrazide. 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Rates of photosynt h e s i s in a t t a c h e d and d e t a c h e d bean l e a v e s , and the e f f e c t of s p r a y i n g with i n d o l e a c e t i c a c i d s o l u t i o n . Plant Physiol. 40:446-451.  121.  Van  Overbeek, J . 1964. Survey of mechanisms of h e r b i c i d e a c t i o n , p. 387-400. J j i L. J . A u d u s , ( e d . ) , P h y s i o l o g y and b i o c h e m i s t r y of h e r b i c i d e s . A c a d e m i c P r e s s , New Y o r k .  109  122.  Vlasyuk,  P.  A.  and  Starchenkov.  I960.  Effect  of  physiologically  a c t i v e s u b s t a n c e s on phosphorus m e t a b o l i s m o f c o r n and s u g a r beet p l a n t s . Primenenie Mikroelementov, Pol imerov i Radiaktivn. Izotopov v S a l ' s k . Khoz., Ukr. Akad. Sel'Skokhoz. Nauk, Tr. Koordinats. Soveshch. 66-73. (C.A. 58:498lb) 123.  Weber, George. 1963. 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E f f e c t of the q u a l i t y of d i f f e r e n t v a r i e t i e s Nauki, Min. S e l ' S k . Khoz. Kaz. (C.A. 63:12248g)  herbicide, 2,4-D, of corn. Vestn. S.S.R. 8:38-40.  on seed Sel'Sk.  APPENDIX  Table and  XVII.  acid  Distribution  insoluble  of  the total  fractions  activity  in control  plants  between  acid  at various  soluble  sampling  t imes  Expt. and Sample ( h r s . )  Leaf Sol. Insol.  Stem Sol. Insol.  Root Sol. Insol.  cpm/g. * * 5 ppm Expt.  II 255 230  8,450  2,320 2,210  360  13,700 11,400  2,550 2,500  2,120 2,050  2,080 3,100  500 235  12,600 7,750  4,150 4,400  262  28  230  28  881  103  16  332  62  185  40  I ,020  210  60  556  99  320  120  1 ,250  270  50  2,080 2,540  4,480 4,420  1,610 1,600  910 910  3,200 3,200  2,130 1,860  70  2,000 2,500  3,415 6,015  1 ,400  890 850  3,000 3,100  1 ,920 1,740  470 340  370  310 160  100 80  820  950  2  14  2,170 2,100  360 595  1 ,770  3,500 4,000  480 730  11,500  60  5 ppm Expt. I l l 0  5 ppm Expt.  ppm cpt. 50  1 ,900  255  8,200  1 ,900 1 ,860  IV  I ,750  V***  70  5 ppm E x p t . VI 1  75  160  580  405  195  270 280  185 110  8,250 10,100  2,230 1,230  3,840 4,650  705 1,020  915  865  8,140 9,620  1 ,330 470 1 ,050 1 ,100  2,360 2,140  112  Table  XVI I .  Expt.  and  Sample  (cont.)  Leaf  (hrs.)  Sol.  Stem  Insol.  Sol.  Root  Insol.  Sol.  Insol.  cpm/g** 50 ppm Expt. I 3  20  48  50 ppm Expt. II 10  20  -> S a m p l i n g exper iment.  2,800  1,870  2,190  4,150  2,870 5,540  2,390 3,380  1,110  1 ,550 2,340  6,340 5,800  4,240 3,280  2,250 2,000  3,730 3,680  2,440  1,710 1,660  times  2,870  7,600  7,630 7,860  8,360 5,820  1,510 1,610  10,250 9,250  6,050 6,100  1 ,325  1 ,710 1,690  865 595  3,800 3,200  3,290 3,180  3,975 4,775  1 ,530 1,390  960 1 ,160  2,680 1 ,960  3,150  correspond  736 858 1 ,220  to hours  after  638 650  spraying  plants  3,875  in each  Counts/minute/gram fresh weight. E x p o s u r e t i m e : 1 h o u r f o r 5 ppm Expt. I l l ; 2 hours f o r a l l other experiments. Holding t i m e : 2 hours f o r 5 ppm E x p t . I I a n d I I I , a n d f o r 50 ppm E x p t . I; 8 h o u r s f o r a l l other experiments. >'w'wV P l a n t s  grown  in complete  nutrient.  113  Table and  XVI I I .  acid  Distribution  insoluble  of  fractions  the total at various  activity times  between  after  acid  spraying  soluble with  5 ppm  2,4-D.  E x p t . and Sample ( h r s . )  Leaf 1nsol. Sol.  Stem Sol. 1nsol.  Root Sol. 1nsol,  cpm/g.* Expt.  1 1  2,280 3,920  425  2,360 2,260  360 300  10,700 10,290  2,960 2,640  6  3,360 5,350  2,020 1 ,300  1 ,920 1 ,910  775 270  7,150 8,350  715 1 ,150  14  2,900 4,570  590 700  2,020 2,150  255 275  12,750 11,400  2,850 2,450  30  5,300 7,320  1,650 1 ,840  3,160 2,690  730 755  10,900 8,700  2,850 3,550  60  2,260 6,600  460 1 ,340  730 1 ,100  2,000 3,400  9,650 11,300  5,100 5,350  2  Expt. 2  11 1  680  416  44  274  36  1,175  145  4  430  60  318  42  1,195  155  8  201  24  210  30  787  123  16  157  33  195  45  1 ,190  240  30  520  70  400  70  990  220  60  128  32  280  40  1,030  -270  50  2,220 1 ,900  4,880 3,060  2,120 1 ,980  770 910  2,840 3,520  2,300 2,550  70  2,250 2,400  4,220  1,750 1,850  1 ,020  3,000 3,400  2,290 2,020  Expt.  IV  3,645  1 ,145  Table  XVI I I.  Expt.  and  Sample  (cont.)  Leaf  (hrs.)  Sol.  Stem  Insol.  Sol.  Root  Insol.  Sol.  Insol.  cpm/g.* Expt.  V** 670  50  215 180  325 190  145 80  1 ,625 770  730 740  425 190  380  190  260  155  835 905  1 ,300  195  9,500 9,900  2,450 2,080  4,720 4,660  1,150 1 ,200  9,750 8,120  3,450 2,760  180 420  70  cpt. 1  * 2  VI  Counts/minute/gram hours  and  920  f o r a l l other  III;  ""<> P l a n t s  8 hours grown  fresh  weight.  experiments.  for a l l other in complete  Exposure Holding  experiments.  nutrient.  time: time:  1 hour  2 hours  f o r E x p t . I I I, for Expts.  II  Table acid  XIX.  Distribution  insoluble  fractions  of  the  total  at  various  activity  times  between  after  acid  spraying  soluble  with  50  and  ppm  2,4-D.  Expt.  and  Sample  Leaf  (hrs.)  Sol.  Stem  Insol.  Sol.  Root  Insol.  cpm/g. 50  Sol.  Insol.  *  ppm  Expt.I  1 ,520  2,070 3,060  565 780  466 660  2,930  7,900  9  3,070 3,080  3,610 2,880  1 ,190 1 ,310  775 630  7,230 7,200  8,800 8,770  20  6,900 5,600  1 ,190 960  1 ,925 1 ,350  875 950  10,150 8,850  4,140 3,960  48  9,500 7,170  1 ,760 1 ,340  2,185 2,460  1 ,970  10,110 10,050  3,130 3,300  2,680 2,490  2,765 1 ,130  1,860 2,090  920  3,310 3,510  3,650 2,850  3,020 5,350  1,675 2,310  1,720 1 ,840  3,250 3,050  3,605 2,750  1 ,440  3  1 ,915  5 0 ppm Expt. II 10  20  "  Counts/minute/gram  Holding  time:  2  hours  fresh for  weight. Expt.  I,  785 1 ,090 860  Exposure 8 hours  time  for  2  Expt.  hours. II.  Table  XX.  ethanol  Distribution soluble  Sample  and  of  the  ethanol  acid  insoluble activity  insoluble  Leaf Sol. Insol.  fractions  ( 5 ppm  Stem Sol. Insol.  between  the  Experiment  IV).  Root Sol. Insol.  cpm/g.* 50  hour Control  Treated  210  700 680  270 280  1 ,860 1 ,580  270 320  320 290  1 ,980 2,260  740  270 600  1 ,650 1 ,140  230 220  2,060 1 ,800  190 270  4,290 4,150  210 180  4,670 2,880  500  145 195  3,270  5,820  150 485  365  150  4,070  650  370  145  3,500  225  920  230  590  1 hour Control  Treated  * Counts/minute/gram fresh Holding time 8 hours.  weight.  Exposure  time  2  hours.  

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