DOCTORAL D I S S E R m m SERIES
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DATE

PUBLICATION NO.

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ANN ARBOR

. MICHIGAN

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This it to MStifg that tht

Th#

B ff« o ts o f th e A a tib io tle
A e ti-d iO B O

M o ry X X is o b o th B o w th o m o

of the tsfisfiBots for
> h.B .

J-y— i,_ a a * H tle 3 r te le s y )

W ilO O B

17 Aug.1951.

o-iti

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m s OTTOLOOICAL EFÏSCTS 07 IHE ANTIBIOTIC ACTI-OIONS

By

Mary X liaab ath Hawthoraa

A D1SSERTÆTI0M

SataBlttad to th # S eh ool o f Qraduat# S ta ll# # o f M lohigan
S ta t# O oUag# o f A g rio o lto r# and A ppllad Solano#
la p a r t ia l fu lflU m a n t of th # raqulrom ont#
fo r th # dogp## o f

DOCTOR 0 7 FHILOSOFHr

D apartnent o f Botany and P la n t P athology

1901

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fsoimspiKtx
FIGUOE
A«

Propha## "raduotlonal'* grouping trc m amtraated mmt#rlal.

B.

Katmpha## "raduotional** grouping from untreated material.

C.

"Scattered** metaphaee with ohroeweome frejpent from root
ti p taken a fte r four hours of eoatlnuoua treatment with
40 ppm solution of Aotl«dlone*

D.

S p lit telophase from root tip taken six hours after
In itia tio n of short treats&ent with S ppm solution.

X.

"scattered** metaphase (c-mltosls) In which chromatids
have "fallen" apart, from root tip taken after nine
hours of eontltmoua treatmsut with 40 pjpi solution.

P.

Disorganised anaphase from root tip taken four hours a fte r
In itia tio n of short minute treatment with 9 ppm solution*

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ACXNOWUOXmDlTS
Th# « r lt# r «Imh## to «zpr### d##p appro#ImtIon to Dr. G. B.
Wiloon a t whoa# anggaatlom th# Inrostlgatloa warn Initiated and nndar
ahoa# diraetion th# projaot aa# conduotad.

Hi# actual counting of a

duplicata aarla# of allda# nada i t poaalbl# to raduca tha aubjactlv#
alanant Imharant In tha method.

Hi# conatant guidance and encourage-

Beat made the completion o f the aork poaalble; hi# patience and good
nature made the aocperlanee an enjoyable one*
To h r. Philip G. Coleman the u rlte r mi aha# to axpraaa her sincere
thank# for a l l of the photographic work and eapeclallj fo r the prepara­
tion of the FlaxlchroBM Frontlapieee which mas Included at his suggest­
ion and which required so much of hi# time to complete.

In addition,

his frequent suggestions during tha course of tha Inrastigation hare.
Indeed, been most helpful.
The w riter Is also Indebted to the following:
Dr. Andrew D. Perejda for the popoparatlon of th# fin a l draft# of
tha text figures.
The All Collage Research Cosomlttee and tha Michigan State College
Hxparimant Station fo r nalrl ng funds e raila b la for tha work.
Tha People of the United State# who, through the G. I . B ill of
Rights, made I t possible f o r the w riter to pursue her graduate studies
of which th is Investigation was an Important u n it.

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Mary UlMbath Hawthorn#
candidat# for

th# d#gr## of

Doctor of Fhiloaophj
Final examination, Anguat 17, 1951, 2:00 P. M., Botany Seminar Room,
Diaaartation:

%# Gytological Effect# of th# Antibiotic Acti-dion#

Outline of Studiea
Major aubject:
Minor aubject:

Botany
2k>ology

Biographical Item#
Born, November 9, 1916, New London, Connecticut
Undergraduate Studiea, Rhode Inland State College, 1953 - 57
Qraduat# Studiea, The Pennaylvania State College, 1938 - 40, %e
Univeraity of Minneaota, 1946 - 49, Michigan
State College, 1949 - 51
Experience:

In#true tor in Botany and Zoology, Undergraduate
Center a Diviaion of the PenneyIrania State
College, 1940 - 44, Aviation Phyalologiat, U. S.
Navy (MR) 1944 - 45, Aaaiatant to the Officer in
Charge of U. 8, Navy Rehabilitation ^braining
School, 1945 - 46, Ina true tor in Botany and
Zoology Uhdergraduate Center a Diviaion of %e
PenneyIvania State College, 1946 —48, Graduate
Teaching Aaaiatant, Department of Natural Scienoea,
Unlveraity of Minneaota, 1946 - 49, Graduate Teach­
ing Aaaiatant, Department of Baaio Biology,
Michigan State College, 1949 - 51.

Member of Phi Kappa Phi, Society of Si(p»a XI, Signa Delta Epailon,
The Oenetiea Society of America, Phi Sigp#a, Tau
Kappa Alpha, Chi Oawga Sorority, American Aaaociation of Thilvaraity Women, U. S. Navy Reaearch
Reaerve Unit 9-16, Lanaing, Michigan.

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tH I CnOLDGIOAL XfTBCTS OP VIE ANTIBIOTIC A0TI-OIOHB

By

Mary Ellaabath &mthcm#

AN ABSTRACT

Sübmlttad to th# Sohool of QroAnoto StuAios of Mlohl^om
Stoto Gollog# of Acrioultor# and Applied Selon##
In pert l e i fn lfl liment of the re q u lre e o te
for the degree of

DOCTOR OF FHILOSOFRr
Deper^ent of Botany end Plant Pathology
Tear

1981

Approved

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Uê t j SllMb«th BÉwthom*
To o tu ij th# nmtixr#, and posaihiUti## of th# origin, of th#
ejtologle#.! #b#rmtlon# produood h j th# antlblotio Aetl-dlon# In th#
m#rl#t#m#tl# tissu## of onion root tip# and to dataxmln# th# thr#ah­
old# of toxle a# wall a# of oytologlcal affaetiranass war# tha main
objaetir## of this inrastigation*
Tha marlatsm# war# traatad with solution# of rarylng ooneantrations ranging from 1 pjm to 00 ppm oontinuoualy for twalra hour#
also for a 10 ninut# pariod foUowad by a d ls tilla d water laaoh# Tram
Faulgan atalnad smaar preparations, a study was s&ada of tha rariou#
types of division figuras which appear ad at hourly Intarrals during
tha f i r s t twalra hours a fte r traatswat was Initiated*
With tha arc option of tha oytologleally Inaffaotlra 1 ppm short
traateant. I t was found that tha f i r s t five hour pariod following tha
in itia tio n of traatsant was tha moat c r itic a l pariod for tha obaarration of tha affects of Actl-diona upon this mariatss&atlc tissue*
Ultimately there was a general trend toward a reduction In tha nwbars
of dirlslon figures following tha short treatment and an arantual
sta b ility In these nusbars in tha oontlnuously traatad root tips*
Dlffaranoas in tha r a la tir a fraquancias of tha indirldual stages of
mitosis appear ad depending upon tha concentration and tha method of
treatment*

Srldanoa was found that tha fraquancias o t tha aberrations

in tha different stages varied; tha highest parcantagas occurred in
pro phase and the lowest in anajdiasa*

Tha frequency parcantagas of tha

sararal type# of aberrations appeared to have bean correlated with time
rather than with either tha concentration or tha method of treatment.

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Mmry E llm b e th Hkwthorm#
No rooover&bl# oberrmtIon# ##r# found.
%# poaslbllltloo of th# lnt«rr#l#tion#hlp of th# ##T#r#l tjp#s
of mb#rration# *# ##U mm th# oytologloal and ganetloal potontialiti##
of each ar# di#eu#s#d.
I t «a# #ugg##t#d that th# thr#ahold of to x le itj may b# vary oloa#
to that n#o#aaary for th# production of th# oytologlcal ab#rratlona
and th# thraahold of #aeh may b# found In aom# combination of oonoan—
tration and tlm# batmoan 1 ppa and 5 pjai and batmaan 0 and IS minutaa.
I t uaa oonoludad that Aotl-dlon# appaarad to hav# acted In a manner
aimllar to that of a aloa fixing agant with a differential affaot upon
tha nuclear and aytoplaamio eonatltuanta of the c e ll and, aa tha dagraa
of tha fixation waa Ineraaaad, tha aararal type# of abarratlona appeared.
Aaaimlng thla analyala of tha obaarvad mitotic breakdown to be correct,
tha following poatulataa concerning tha a c tiv ltlaa in normal mitoaia
were offered:

(a) a gradual formation of tha aplndla, (b) cytoplaamic

origin of, a t laaat, m«mm part# of tha apindla, (e) tha poaaibility of
tha origin of anaphaaa monreawnta from aoaie machaul ama arlalng within
tha chromatid# themaalvea.

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Mary Slisab«th Bmwthcm#
Selected Blbliogrephj
Allen, N. S., G. B, Wlleon end S« Poeell (1990)
CoBperetire effects of colchicine end sodium nucleate on scmetlc
chroBoacmss of AHiqn end Tfedesoantie.
Jour. Hsred. 41;159»Î63.
D*Aaeto, F. (1949)
Preprophase inhibition of mi tor is in root merlst«BS.
Ceryologie 1:109-121.
Gelinsky» I . (1948)
The effect of certain phosphates on mitosis.
Genetics 38:610-611.
Haskins, C. L. (1948)
Segreeeticn and reduction in soBstic tissu es.
HUB ce
cena.
tions in Allium
Jour. Hsred.r8?:5ll=3)
39:811-388.

I . I n i t i a l observa­

Kodani, M. (1948)
Sodium ribose nucleate and mitosis. Induction of morphological
changes in the chrcmoscmes and of abnormalities in mitotic
divisions in the root meristems.
Levan, A. (1938)
The effect of colchicine on root mitosis in Allium.
Hérédités 24:471-486.
and T. Lotfr, (1949)
Haphthalene acetic acid In the Allim* te s t.
Hérédités 35:337-373.
Ostergren, F. (1944)
Colchicine mitosis, chromosome contraction, narcosis and protein
chain folding.
Hérédités 50:429-467.
Made, B. (1990)
The mechanism of mitosis baaed on studies of the submicroscopic
structure of the living state of Tradescantia c e ll.
Cytologie 16:1-26.
Wilson, G. B. (1950)
Cytological effects of some antibiotics.
Jour. Hered. 41:226—
231.

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TÀBIZ or CONTENTS

TAOX
INTRODUCTION......................................................................................................................

1

UÀTXRIALS AND lORHCDS.................................................................................................

IN

OBSERVATIONS

21

.................................................................................................................

Qualitative Aapaota • • • • • • • • • • • • • • • • • • • •

21

Quantitative Aepaeta

94

..........................

.

. .

. . .

General Effeete of A otl-dlone............................................
Specific Iffe e te of Aetl-41om e.....................

34

. . .

. . .

DISCUSSION...........................................................................................................................
General Iffeete of Aetl-dlone .
Specific Xffeeta of Aetl-dlone

.

.

.

.

.

.

.

.

.

.

. . . . . . . . . . . . . .

49
69
69
67

The Typee of Aberratlone, th eir Interrelatlonehlp and
Signifieancea . . . . . . . . . . . . . . . . . . .

67

The Cytological Aberrations, their Relative frequencies
and Relationships to ^rpes of treatment . . . . . . . .

73

SONNAKT.....................................................................................................................................

83

TEXT FIGURES

87

. . . . . . . . . . . . . . . . . . . . . . . .

PLATES.....................................................................................................................................

97

......................................................................................................................

103

......................................................................................................................

116

APPENDIX I I ........................................................................................................................

117

APPENDIX I

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1%ST 07 lABIZS

TEXT TABUS
I.

PAGX

Rooting Media end Subeeqnent Areetment# of the
Terlou# Serlee • • • • • • • • • • • • • • • • • • •

II.

Senple Copy of the Date Sheets

III#

15

.....................................

19

Average Member of Division Figures Bar Axis Obtained
from the Combined Data of F irst Five Hours • • • • «

IV.

36

Peroantages of Aberrant Figures Obtained from Combined
Data of F irst Five Hours . . . . . . . . . . . . # .

V.

60

Mean Peroantages of Prophaee and Metaphase Cells
Containing "Reduotlonal** and "Segragatlonal"
Chrouplngs

VI.

. . . . . . . .

•

68

Cytologloal and Ganetloal P o te n tia litie s of the
Several ^rpes of Aberrations . . .

69

APPENDIX TABUS
APPENDIX I
I.

Reported Results of %eaMsents of Merlstssmtlo Tissues
with Organic Chemicals . . . . . .

II.

. . . . • • • • •

116

Reported Results of Treatments of Merlstsmatlo Tissues
with Inorganic Chemicals

III.

........................

U6G

Reported Results of Application of Certain Physical
Stimuli to Merlstmmatlc Tissue *

.............................

116H

APPENDIX I I
1 - 19C

Percentages of Total Division Figures of Individual
S ta g e s...........................................................................................................117-139

16 -

28CAverage Numbers of Division Figures Per Axis .

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• . • • 140 —146

IXST OF T4BUBS (oontlxna«d)
AFPSNDIX II
£9 - 32

PA
Q
EI
Hourly Change# in Average Ntniher of Division Flgir##
Per A x is

147 - 148

35 -

47C Pereentage# of Aberrant Prophase#

48 -

62C Percentages of Scattered Metaphaee#

69 -

770 Percentages of Disorganised and Total Anaphases

78 -

900 Percentages of Cells Containing "Reduotlonal" or
^Segregatlonal" Cferouplnga

.

.

• • • • • • • • •

149 - 156

• • • • • • • •

157 - 164

. . .

. .

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• #

165 - 177

. .

178 —184

LIST OF ISXT FIOORXS
TEXT FIGURES
1#

PAGB

Average Amber# of Divlelom Figures Per Axis Obteined
from Combined Date of F irs t 5 Hours • • • • • • • • •

2 - 13#

Hourly Changes in Average Amber of Division Figures
Per A x i s ......................................................................................................

14 - 19.

........................................

. .

90

. . . .

. . . .

................................

. . . . . . . . .

94

Pereentages of Aberrant Figures Obtained from Combined
Data of F irst Five Hours

73.

93

Hourly Pareentages of Disorganised Anaphases Compared
with Pareentages of Total Anaphases

70 - 72.

92

Hourly Pareentages of Aberrant Hetaphases Compared with
Pareentages of Total Hetaphase. . . . . . . . . . . .

56 - 69.

91

Hourly Pareentages of Aberrant Prophase Compared with
Peroentagss of Total Prophase

42 - 55.

89

Hourly Variation in irequanoies of Individual Stages
in the Intermittent TTealsmnt

28 - 41.

• • • • • • • • • • • •

Hourly Variation in Frequensies of Individual Stages
in Continuous Treatment

25 - 27.

88

Hourly Fhriation in Fïrequanoies of Individual Stages
Following the Short Treatment

20 - 24.

87

95

Mean Pareentages of Arophase and Metaphaee Calls
Containing **Segregational** or "Reduetional" Groupings
Found During F irst Five Hours

. . . . . . . . . . . .

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96

LIST OF FZATSS

FACE
FBONTISFm

Ml#e#U#n#oo# droop of Aborront Flgoro#

FIATE I .

**lionMlIy** OrganlMd Flgur##

97

FIATB I I .

A Sorlos of *^odtMtlonol" d ro u p ln g # .....................

98

PIATB I I I .

A Sorioo of *'8#gr#emtlonal" Grouping#

99

FIATB IV.

A Sorio# of **Seatt#rod’* Figaro# of the
Diaorgonizod "%rp#

FIAIS V.

.

.

.

.

.

.

.

• • •

. . . .

.

.

.

A Sorio# of Coll# Shooing Indication# of
Toxicity ..................................... . . . . . . . . . .

FIATS VI.

100

MiocoUanoou#

.............................................................

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101
108

IMTSODUOTIOK
Th# o#U mmj b# eoa#lA#r#A # pivot around «hloh tb# ftowtioB#
of living oreanimm# ravolva.

Whan oonaldarad in thia light# tha

a e tiv itiaa of tha living oall aaaima auoh fundaoaantal aignifioanoa
that ona ia atimmlatad to aaak anaaara to naaolvad biologioal problana
a t tha oallular laval*
Ona of tha intriguing proeaaaaa of oall# that# ao far# ha#
dafiad anolyal# i# tha aaahanl wm of oall division*

IWon obaarvation#

of tha prooaaa and atudio# of f izad n otarial oontalnlng dividing oall#
thara hava avolvad nany hypothaaaa but fan unoontaatabla thaoriaa aonoorning tha aaohaniaaa involvad*

Sinoa tha atndj of anonalia# ha# boon

ao f ru itfu l in othar branehaa of biology thla taohniqua naa adoptad by
oytologiot# aa a jronioing routa tenard a battar undarstanding of thaaa
parplaxlng maohaniona.

Aooordingly# dividing oall# hava baaa subjaatad

to alnoot ovary oonoaivabla typa of ohonloal and phyaioal traateant and
tha affaot# notad*

I t ia tha baaio asomption in th is raaaarah that by

uatohing tha prooaaa braak doom ona aan# parhaps# find aena of tha Bias­
ing olnao to tha naohaulm# althar in tarn# of foroaa or grooaaaa# in­
volved, or in term# of a battar understand 1ng of tha n ativ ities of the
ohrcnosanos thomaalvaa.

Innuaarabla invoatigatora hava purauad this

tack and the lite ra tu re resulting from their labors is besoming inoraaaIngly extansiva*

A oomplata ravian of th is previous nark is beyond tha

soopa of tha present report, but such an activ ity nould undoubtedly
resu lt in an assemblage of facts nhioh# nhan placed side by side# m i^ t

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2

fo rm th# foundation of an entlraly nuw hjpoth##i#«
MOT purpo8#8 of tha orientation of th# presant raaaarah, a
m a m m rj of aona of thaaa invaatlgatioo# aaa praparad and will ba found
in Appandix I#

Skma of tha rafaranoaa oitad thara oarry axtanaira

bibliographiaa which ahould ba atodiad bafora tha oonplata pietura
will ba aaan.
Tb# unaroidabl# aubJaotiTity that paznaataa thia lita ra tu ra ,
oouplad with a eonplat# lack of atandardlsation of taxma, maka tha
of tha praparation of auoh a tabla a moat d iffic u lt ona.

An obaarmticn

which ona inraatigator may conaidar to ba paramount in algnlficanoa an­
other may hava naglactad or orarlookad antiraly.

In aona caaaa i t appaara

that tha aama condition haa been given different taxma.

In thia table

notâtiona ware made only in thoaa caaaa where apaeific atatamanta war#
made concerning tha particular condition,

Thara ia a more or laaa ganaral

agraanant that tha affecta produced are corralatad with tha comaantra tion
uaad,

Tha table reporta tha cytological affecta throughout tha aariaa of

concentrât iona uaad.
Tor purpoaaa of thia diacuaaion aoaw ganaraliaationa may ba draan
from tha accimiulatad reporta,

Tha phaaaa of thia material that lanadi-

ataly impinge upon tha praaant reaearch may ba divided into too par ta .
1.

Tha ganaral raaulta obtained from othar typaa of treatment,

2,

Tha conaanaua of opiniona concerning tha nature of tha mach-

aniaa involved in tha braakdowna,
Thaaa divaraa treatment a, both chemical and phyaical, produce
raaulta that may ba claaaifiad into two overlapping aatagoriaa:
mitotic diatnrbancaa; and (£} tha chroa&oaoaial abarratlona.

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(1) the

3
On th# b*#l# of th# ultlaat# #ff#ot# prodoo#d th# mitotlo dimtorb#no#s mmj b# furth#r p#rtition#d into two mub-sroupm: (#) Thom#
which can b# porpotwmtod mine# th# e#Um g#n#mliy r#oor#r and contlxroa
to dlTld# according to a nor# or l#ma regular pattern producing daudhtar
call# which carry on th# new condition; (b) %oa# mitotic dimturbancam
the affecta of which, a t praaant, hav# not beam parpatuatad mine# tha
call# ganarally do not racovar from tha diaorgarni aation induced by tha
treatment*

Tha f i r at group would contain tha polyploidiaing agent# much

aa: colchicine (18, 40, 48, 70 and 99), eye Lochlorohaxana (33, 7 t, 104
and 115), varatrine (144), acanaphthana (70, 79 and 99), and chloral
hydrate (95 and 101)*

Tha aacond group would contain thoaa aubatanoaa

which are potentially capable of producing polyploidy, of inducing raductiona in chroaioacma nuabara and, coaaaqwantly, raaulting in ganatical
aagragatiom#

Thaaa potential polyploidiaing aganta raault in tha condi­

tion that haa now baccaia known aa "c^mitoaia*"^

In tha abaanca of tha

aplndla, tha chronoaomaa are acattarad throughout tha c a ll and tha dalayad
diviaion of tha kinatoohora raaulta in tha production of I-aha pad chrcmoaomaa*

Eventually tha chromatid# may f a ll apart and raatitution nuclei

may ba formed but, aa pravioualy mentioned, thaaa call# ganarally do not
divide again*

Thna they arc only potentially capable of producing

^ "C^aitoaia" ia tha term auggaatad by Levan (76) to refer to tha
type of mitoaia produced by eolchicina. In thia raapact tha term ^
impliaa tha polyploid raault* Sinoa tha appearance of calla follow­
ing other typaa of treatment# ia practically a duplicate of that
following colchicine but (aa pointed out above) thaaa conditiona do
not nacaaaarily lead to polyploidy, tha t e n haa now coma to refer
to the two outatanding affecta of colchicine treatment, namely,
aplndla di eruption accompanied by a delayed aplittimg of tha kinatachoraa of tha over-contracted chromoaomaa*

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4
polyploidy*

T&blo# I mad H of Appmadlx I mho# tho dirormifiod typos of

ohmmlomlm that tamcro horns roportod m
m ompmblo of produo log thim motion*
SoBo of thomo mrox phomphmtom (08), nitrmtom (88), UDT (105), mntibiotiom
(148), mmltm of mmoloio moid (69), othylono glyool (107), tolmidino blno
(8), oortmla monooyolio orgmaio oompommdm (86), mad oortmia amphtholoao
dofivmtivom (86)*
Am prmvioumly moatioaod tho tromtaontm #hieh may rommlt ia s ito tio
ooaditioam mhioh potoatimlly mmy lomd to tho roduotion ia ohroBomcmM am&borm oad,,monmoquoatly, gmaotioml mogrogmtioa, mhould bo plmood ia thim
Boii*roeorormblo omtmgory*

Tho tormm "redmotionmJ." or "mmgrogmtioaml"

dfoupiagm oro fro%mootly mood to rofor to thim mitomtioa (1, 63, 64, 110,
148 ot ml).

Boomumo, mt f i r s t ho fo lt ho hod found ma iMBOlOdJ #ith tho

rogulmr mothod of rodumtioa prooodlag tho mhmago from nporophyto to gmaotophyto ammborm, Bomkiam origiamlly umod tho toxm "mommtio moiomim" to
rofor to thim mitumtioa (63).

Thim typo of offoot im roportod m
m muoh

mftor tho followinc tromtmoatm: oodimm auolomto (63), oolohioiao (8), mad
Aoti-diono (148)*

Tho "oxplodod oHmitomom** of hmrbor mad Omllma (6) mad

tho **dimtributlTo o^&itomom" domoribod by Bfbom mad Kautmmom (106) mppomr
to iavolvo tho momo mrrmaeemmnt mm that torood mboro "roduotioaml" or
"mogrogmtioaml" groupiagm*

Oa thim bmmim momio ohmmlomlm mmy bo mddod to

tho l i s t of tromtmwats that inorommm tho iaoidonoo of thomo mrrmagmaoatm
of ohrooomcmiom ia motivoly dividiag timmuo; muoh m
mmothyl amphthOQuiaoao
(106) mad tho aitrmtos (88)*

I t mhould bo poiatod out horo that boomumo

of thim di f f oromoo of opiaioo oonooraiag thoir mignifiomnoo i t im guito
likoly that thim offoot ham boon ororlookod by momo iavomtigmtorm, mad

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s
that many aora ahaml oala may ba fovnd to eauaa tha ■— affaat i f latar
morkara ara alartad to look for tham#
Among tha ohromoaoma irragulmritiaa ia A.ppandlx I "atlaklmaaa,"
ahortanlmg, laagthanlng* aroai<m» and fra^pmaatatloma ara llatad*

Of

thaaa» probably tha aroalooa and fra#mamtatloaa ara of graataat aytologioal and gaaatioal aignifioanoa.

Ona la impraaaad with tha almllarlty

batwaan tha ohamlaal and phyaioal traatmamta la prodnoing aoma of thaaa
affaata on tha ohromoaoa&aa.

Rpa#p»antad ohromoaomaa hara baaa raportad

a fta r traatmamta with tha following ohomleala: aoanmphthana» 9 amlnoaorldlna and phoaphana-5-O (34)» mathylana and tolmidlna bloa (8),
bomaol (43)» mitrogam mnatard (105)» radlophoaphowona (44)» and aararal
phanola (8t and 88).

Tha aoma raault haa baaa obtalnad through tha naa

of tha following phyaioal aganta: I^raya (87 and 180)» faat aantrona
(57)» and ultraaonlo vibration# (108).

I t may ba of Intaraat to nota

hara that Lovan and TJlo (87) polmtad out that» nnllka X^aya» tha
phanola aaldcm indnoa traaaloeatlon and raportad that tha damaga oanaad
by tha ohmtienl la outtlng through tha ohromoaoma th raad uaually doaa not
allow hanllng.

Vbathar or not thla la trua of othar olMmloala aould not

ba dlaoamad from tha raporta.
Amarona Inraatigator a hava attomptad axplanatlona of tha moohant m
of o-mitoaia.

Moat of tha work waa baaad upon tha aaavmptloa that» baaanaa

of tha almllarlty la tha appaaraneaa of tha raault» maahanlam waa aimllar
for a l l traaWanta and oommon faator In a l l traahaanta waa tha ma in objaotIra In thaaa raaaarohaa.
propoaad.

Both ohamlaal and phyaioal axplaaatloma hava baan

On tha ohamlaal aida» tha aarly workara attamptad to laara idiloh

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6
p#rt of tho eolehieiao Boloeulo ooo rooponoihlo for tho o e tiT itj.

Ao o

rooult of th o ir oork, Bmoo oad Cohon (80) poiatod out th a t, of tho
Toriotj of aoloenlor grouping# uhieb ocmpooo tho oolohioiao molooulo,
i t #o# not poBBiblo to Bay th at any singlo group in ooaontlal for tho
aotiTity of oolohioiao on mitoBio and booauoo of thia laok of Bpoeifioity
urgod an orami nation of othar aynthotio oonpoundo of analogoua atruoturo.
Polloaing thia aork, a largo oorioa of orgaaio oompounda ana toatod and a
groat aany roporta appoarod.

A rorioa of thoao oarlior opinion# aaa pro-

aontod by Lovan and Oatorgroa (66)#

Aroa tho la tto r aurroy ono nay doduoo

that tho divaraifiod ocaoluaiona aoro about aa nnaoroua aa tho iavoatigatora, and tho ao tlv ity aaa attrlbutod to ahatovor particular variablo tho
aorkar happonod to ohooao to vary.

T o r oxanplo, tho a c tiv ity aaa aaoribod

to tho aroantio nuoloua by Siaonot and Ouinoohot (187), to tho —Ino groupingo by Favoraky (47) and to aca&o ohomioal atruoturo a in ila r to th at of tho
oaroinogonlo hydrooarbona by Shauok and Ouaaova (185).

Finally, Stoinoggor

and Lovan (153), aorking a ith iao-oolohioino oonoludod th a t tho "o^ing of
tho oolohloino molooulo la of doelaivo aignifioanoa for tho apaoiflo activ­
ity on Alliua roota, booauao i f tha koto- and aothaxyl-groupa of ring C
aro Intorohangod, aa i t la in tho oaao of iao-eolohioino, tho apooifio
offoot of oolohloino diaappoara.**
A foa axplanatlona atom from tho baaio aaammption that a phyaioal
moohaniam la involved in tho production of o^&ltotlo a c tiv ity .

Althou^

ho aaauaod aoma ohangoa of a more "qualitativo" naturo to bo proaont,
Wada (159) oplnod th at tho action of oolohloino ia mai nly duo to it#
a b ility to roduoo tho aurfaoo ton#ion of tho "atrakopla»a" and 6hlgonaga
(121) oonolWod that tho apindla dlaturbanooa produced by tho varioua

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T
poisons» ehlom l harArnts» nleotlns, sud onffeins, opsmted in %msnnsr
similmr to tbnt of hypertonie sngsr solution» thmt is» they esused #
dehydretiOB of the Icln o p lei* (spindle or i^urs#^plsst)#

I t might be

of in terest to nets that both of these l e t te r Ideas mere developed from
observations made oa living eelle*

As a great variety of ehsmleals of

midely divergent nature mere reported in the lite r a tu re as eapable of
indueing e«eiltoeis» Levan and Ostergren (8d) noted th at the vast najority
of these materials» mhioh mere meet active» mere least solnble in mater»
and began vorldag on a nom hypothesis*

la testing the alpha and beta

derivatives of naphthalene they found a defin ite positive, inverse eorrelation betmeen mater solu bility and the thresholds of e««itotio notion;
i.e*» the la t t e r increased as the former decreased*

This fast» eoupled

mith the vide divergence in the chemical nature of the c-mitotic sub­
stances» as ms11 as the fact that many of the narcotics tested mere found
hi^ily effective» led them to the opinion that the eesential nature of the
c-mitotic a c tiv ity is physical and does not depend directly upon the chem­
ical composition of the substance.

Because of the inverse correlation»

they arrived a t the comelusion that fundementally the acticn of these
e^&ltotic eubstanoes mae a nareotie one and that the "Meyer-Overtom theory
of narcosis may hold in the cases of ths c-mitotie substances» i.e .» the
decieive concentration of the substance may be that ia the lipoids» not in
the mater phase of the cells*"

Ostergren (100) elaborated upon th is idea

and described the protein chain folding he believed responeible for
spindle destruction a fte r the penetration of the subetance*

Be recognised

that colchicine does not f i t into th is hypotlMSis and because of th is sug­
gested that some substances probably react chemically*

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8
Th# ob##rv#tlomm that #cm# of th### #h#mle#l#, in addition to In­
hibiting th# #plndl#, had othar affaot# atartad mom# n#« linam of think­
ing.

rodani (69) not ad, among th# varlon# affaot# of aodlia nnalaat#

aolutlona, an inhibition of ohroakoaom# radupUoatlon and th# formation
of aklnatle frog^anta.

D'Amato (81), a fta r taatlng a aariaa of ahamloala,

obaarvod that tmo dlffarant affaot# mnat ba dlatlngoiahad:

(1) prapraphaaa

Inhibition and (8) an inhibition of apindla formation giviag riaa to omitoala.

Ha maa of tha opinion that "tha aplndla-inhibiting affaot, mani-

faatad in tjp ia a l qmo&itotla polaona, may ba of qnita dlffarant aort than
that by mhloh prapraphaaa polaona Indnaa tha OHoitotla affaot."

Oalinaky

(98) found that fellomlng tha phoaphata traahaanta tha dariatlona from
normality mara obaarrad during pro^&aaa in tha dalay of nnolaar mambrana
"braakdomn."

Ha polntad out that th ia affaot had not baaa obaarrad aftar

tha uaa of tha o^âltotlo polaona.

Thaaa aubatanoaa had baan raportad to

parait mitoaia to prooaad normally (76) u n til mataphaao mhan thoir apaoiflo
offoot on tha aplndla maehaniam maa manifaatod.

Aooordingly, ha oonoludad

that, "tha nnoriantad promotaphaaa ohromoaomaa mnat tharofora larolra
foroaa othar than thoaa aaaoolatad mlth tha oriantod apindla maohanimm."
A diatlnotlon batmoan tha ^mitotlo polaona" of uhioh oolahiolaa maa aonaldarad a typa aubatanea and tha "phyalologleal aubatanoaa" axomplifiad
by tha nuelaata aalta maa mada by Allan, Wllaon and Pomall (8).

Working

from tha ehamloal rlampolnt, Loralaaa and Rarall (91) attomptad to oorralata tha ohomlatry and modaa of notion of oortain of tha dlffarant alaaaaa
of oompounda th at had baan daaoribod aa baring an affaot upoa dlrlAlng
oalla.

Thay f a i t that tha "mitotlo polaona" eould ba elaaalflad Into at

laaat thraa dlatlnot alaaaaa aaoh of mhloh mould aham a ohamlaal
a

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9
homogeneity eorreleted with biologioel eetlvlty:

(1) eompoonde known for

eome phyeiologienl eetlv lty whleh Include the beete r loetet le end entlprotoaoel egente end entl«growth eubeteneee of plent origin which exert
e highly toxic, nom-epeclflc effect upon the c e ll, (fi) the veeleente,
eueh ee eulfur, nitrogen nmeterde, dl-epoxide# end preteememomln, which
here e highly specific effect upon the resting nucleus which is menlfested in subsequent mitosis by the eppeerence of breekege end reerrenge*
neots of verious kinds, (S) the eompounds cepeble of destroying or pre­
venting the foxnetioa of the mitotic spindle without undue toxic effect
upon the c e ll, such es colchicine.

Hence, the question of the underlying

mecheni cm of cHcltosis remelns unenswer ed, end w ill probebly continue to
be unsettled u n til the

underlying Issues relmted to the modes of opere-

tion of the severel treetments ere f i r s t decided upon.
The objective of these previous investlgetions wes primerily the
determinetion of the type of effect produced by e perticuler chcmicel,
the chemicel group, or redicel responsible for the reectiom.
"effective" concentr e t ion threshold wes stressed.

The

The technique for

determining th is threshold essentielly involved the eppUcetion of the
chemicel for en esteblished period of time.
getors the time period verled.

Among the severel investi-

T o r exemple, four hours, twelve hours

end three deys were the esteblished times for exeminetion of test neteriel
et the Cytogenic Leboretory e t Svelof (151).

This wes the stenderd pro­

cedure for Leven, Ostergren, Steinegger end e l l workers et thet stetion.
The Wisconsin group mention exsminetion of th eir meteriel et "verious
times" (SS end 69).

Hence, the bulk of the present litereture conteins

reports of obeervetions mode on fixed meteriel efter continuous treetment

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10
with diver#e ehwaleale e t verylag eoneeatretloae eed e v in e d after
rather wide iatervala of time.
Some iaveatigatora have been studjing the effeete of otaemioal
treatment# on living tisane (100, 121 and 159).

In th is reaearoh,

stamen hairs or yonng petal o ells, from the hods of Tradeseaatia sp.,
provided the dividing o ells for the work.

Althongh these experiments

have resulted in some very interesting observations, in using th is
technique, the worker is limited to the reaetions of a single c e ll at
any particular time and, of course, is encumbered by optical limita­
tions when waking deductions from unstained c e lls .
This is a report of a detailed quantitative analysis of the cytologioal effects of a single chemical, the a n tib io tic, Actl-41oae.

In

a preliminary report of observations made a fte r 24 hours exposure to
100 parts per million, Wilson (142) showed that the effects of th is
substance run the gamut of aberrations produced by most other chemicals.
Accordingly, he stated that th is "is one of the most interesting anti­
biotics yet investigated since I t r eadily and comsistently induces spe­
c ific aberrations in sdtotlc behavior."

for th is reason i t was sales ted

for th is study.
By varying not only the concentration, but also the time of tre at­
ment, and by studying the effects produced in eequential periods of tine,
particularly in the c r itic a l early interval a fte r exposure, i t was hoped
that data could be obtained that would give some specific clues concern­
ing the origin of some of these aberrations, and would yield some infor­
mation concerning the nature of, i f not the neuhsiilcn involved in, these

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11
lrr#galmrltle#.

Mar# •p#cirioallj» by # quoatitatlT# atuAy of th#

#b#rr#tloii# i t ### hoped that I t mould bo pooalbl# to loom ^Aothor or
not any apooifio rolatlooahlp ozlotod b#t##on tho ooourrono# of th# I r rogularltl## and aueh faotora a# th# oomoontratlon, th# tlm# and th#
mothod of tr#atm#nt.

With th### data arailabl# I t #a# thought that i t

oottld b# poaaibl# to dafino, in term# of eonoontratlon and time, a par­
tic u la r thr#ahold at mhioh recoverable aberration# might be obtained.

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MASERIAIS AND KXTBODS
Th# Amgloiaml aBtibiotio Aoti-dlon# (197 and 141) jorodoo^d by
8tr#*ta#ye## mrleeu# (49 and 79) and aaad la this aarlaa of iaraaticatloa#, aaa obtaiaad throogh th# ooart##y of th# Dpjaha Oompaay,
Kalaeasoo, Mlahlgaa.

With a malt lag pelat of 119 - 118^. aad a oolm-

b l l i ^ of 11 ag./ml. at 1^0. aod botmaoa 89 to 90 ag./al* a t room tomporataro. I t ha# a ehomloal formal# a# foUomo:

e iv -o t

Th# root tip# d#T#lop#d from th# eo#a»oa oommarolal joUom variety
of Alll«m C#pa 1. iroTld#d th# m#rl#tom#tlo tl##a# for th# #tady of th#
oytologleal offoot#.

for praotleally all of th# #zp#rla#at# la#lad#d la

thl# report, th# bulb# mar# obtained from a alagl# orop prodaoad by Nr.
Wall Poataofc of Xl#l#, Mlehlgaa.^
Thro# ##rl## 91, 95 aad 95 had boon eomplotod boforo th# Poataak
onion# mar# obtained but thl# #upply aa# axhaaatod in #ub##qa#nt teat#
no i t oa# not poaolbl# to daplloato them. Homovor# alaoo dapUoato#
of those throe sarlo# produood ooaolstent result# I t mas #om#ld«red
unaeoessary to repeat these treatments.

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IS
la th# ##rll#r ##ri## th# bulb# ##r# root#d Im tap uat#r maA th#
tr##tm#at solution# u#r# ir#p#r#â with d i# till# d uator.

Aftor tre a t­

ment aad a thorough uaahlag with diotiU ad water, a l l oaioa# war# them
plae#d la d la tlU a d water uhleh was ebaaged every hour for the roeorory
te eta.

Later, th# oal oa# b#gaa to produe# fewer aad fewer root# la the

tap water prtrrlded la th# laboratory aad I t b a o ie meeeeeery to ohaag#
thl# teehaiqu# #ow#what.

After teetiag water from a variety of eoureee.

I t wa# fouad th at aerated d le tllle d water wa# th# meet eueeeeefal root­
ing medium.
se rie s.

Aeeordlagly, thl# proeedur# wa# used for a l l the subsequent

Th# rootlag medium aad th# treatmeat th at followed 1# sheua la

Table X.
TABU I
ROOTINB MEDIA AM
D SUBSEQUENT BtEAlMENTS OT THE 7ASI0D8 SERIES

TAP W
ATER

ASATSD DISmiED EATER

Series

Series
PPM

98
96 h 180
118 h 128
188
95 & 188
108 & 115
186
187

80
40
80
80
40
80
40

Duration

Short (15 mla.)
Short (15 mla.)
Short (15 mla.)
Oontlauou#
Contlauou#
Oontlaueus
latezm ltteat
(15 mla./8 h rs.)

FfM

Treatmeat
Duration

149 A 174A
128 & 158B
140

1
5
80

Short (18 mln.)
Short (18 mla.)
Short (15 mla.)

146 A 181
159 A 158A
124

1
5
80

Oontlaueus
Oontlauou#
Oontlaueus

135

40

latezmltteat
(15 mln./I hr.)

untreated
138
(Coaeeeutive hre. )
1378

untreated
(Coaeeeutive hrs.)
untreated (Seme hr.!

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14
HaoltelAM «ad bm r

eO"^#orh#r# (fS)

#%adl#é th#

•ovro# OB th# toocieltr of #om# »#reurl#l poi#ea#«

#fr#ot#

of ##t#r

Th#j r#port#d th#t

th# tr#a#f#r from t#p to dlotiUod m#t«r r#t#rd#d growth of th# omiem
root tip# mad prorod f a ta l momatlm## mal### th# ohaag# «aa mad# threogh
mixtar## of th# two «atarm#

la addltlom, thay poiatad oat that almoat

aaj amddam phyaloal ohaag# la th# madlwa Impadaa growth.
la nwarna# toat# ooadaotad la th la laboratory a l l aorta of «atar
•cmblaatioaa war# tria d la th# gaaaral proeadar# pattara of rootlag,
traatlag aad laaehlag a fta r tra ata a a t.

So fa r, aa appraoiabla d lffa r-

aaoo# haw# baaa obsarrad aanng tha a*aroue oomblaatioaa laraatlgatad.
I t anat ba poiatad out, howaror, th a t la thaaa atudlaa a traaafar from
althar tap watar or traatmaat aolutloa dlraotly to thoroughly dlatlU ad
watar waa mot mad# aad hamaa, tha poaalblUty of tha phyaloal ahaak of
auddaa ohaaga la watar aloma earn ba a lla inatad.

la addltlom, tha dla-

tlU a d watar aval labia la thla laboratory la omaa dlatlU ad la a matal
a tm

aad la mot aomplataly fra# of a l l loaa.
Slaea tha roota tolaratad mora polaoa la "raw** watar takam bafora

purlfloatlon la tha watar worka, kaalhrlaaa aoaaludad that tha Imbalaaaa
of loaa la o lty watar playa aa Importaat rola la dablU tatlag tha tlaauaa
aad urgad that tha toalolty affaata ahould ba tafcaa lato aooaldaratlom
la tha la ta ra a t of tha ataadardlwat 1om of tha AUlma ta a t.
I t w ill ba Obaarrad la Thbla I th a t, la thla laraatlgatloa, for
thoaa aarlaa la whlah tha graataat dllutioaa war# taatad, oaioaa wara
uaad which had baaa rootad la aaratad dlatlU ad watar.
aoaeaatratloa thara waa oaa aarlaa of aaah typa.

For tha SO p ^

Eafaraaaa to Appaadlz

I I , Tablaa 9 ami 8, w ill raraal that tha raaulta obtaiaad from tha oaioma

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IS
T ootm à la tap aatar aar# ooaalataat with data obtaiaad tr c m thoaa rootad
la tha d iatiU ad a a ta r.

la tha hlghar aonoaatrat Ica aarlaa tha traaafera

aara mada frcm tap aatar to 40 aad 80 ppa aolatloaa.

Oaa ooatrol aarlaa

for aaah rootlag madl«m la laaladad la thaaa ra a a lta .

la tha hlghaat

diXatlona (1 pp& aad 5 pf&), rootlag, traatmaat aad raaorary aolatloaa
aara proparad from d latlU ad aatar thna praoladlng tha poaalbiU tj of a
d latortlo a of ra aa lta b j tomlalty affaota.
A moro or laaa ataadard aathod of proaodnra fo r tha traatmamta aaa
aaad# VOr rootlag, tha bolba aara arramgad om aaalT glaaa oomtalaara la
aaah a aannor th a t from oaa-fearth to omo-half laoh of tha balb pro tr udad
lato tha rootlag modlm&* Vhaa roota aara from oma aad oma^half to tao aad
oao-half iaohaa la laagth tha root tlpa aaro oandaad aad. I f h l ^ mltotlo
a o tlT ltj aara la arldanoo, traatmaata aaro atartod by plaaiag tha young
roota la tha traataaat aolutloa aad auppor t lag tha bulb orar tha aoatalaor.

Whoa traatmaata aara oomplatad a l l bulba aar o plaoad orar dlatlU ad

aator ahloh aaa ohaagad ovary hour aad a ohaok for rooovary of tha
tlaauaa aaa mada.
Root tlpa aara oollootad la a throa to oma abooluto aloohol-aoatlo
•old mlxturo, flzad fo r flfto a a miautaa a t dO^, hydroUmad la oaa mormal
hydroohlorla aold for oight to ton miautaa, atalaad with faulgaa'a atala
•ad amaar praparatioma mada.

Aftar dahydratlom la a 90# athyl al aohol

to ahloh a U t t l a faat graaa for oouataratala had baaa addad, tha alldaa
thaa aara mada parmaaaat a lth dlaphana.
A atook aolutiom of 100 ppm aaa praparad by dlaaolviag tha poadarai
Aotl-dioa# la d latlU ad aatar aad kapt rofrlgaratad u n til tlma for uaa.
Whoa dllutioaa from tha atook aolutiom aara praparad, aara aaa takaa that

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16
th# treatment eolation# ###t#med room temperatare before the root# mere
plaoed la them.
A preliminary aerie# of experiment# mere oooAuoted In alileh th#
root tip# mere ooUeoted every fifte en mlnate# and examined.

Count#

made on the## elide# indicated that thi# interval aa# too ehort to re­
veal change# ocearring and i t aaa decided that daring th# coarae of th#
farth#r eerie# th# root tip# would be eoUeoted every hoar.

I t aa#

farther decided, a# a reault of the## exploratory \##t#, that tao main
type# of treatment# mould be foUomed for the detailed, comparative,
quantitative studjt

(1) a abort treatment In ahloh th# young root# mould

be axpoeed to th# te a t eolation for a period of fifteen minute# and then
tranaferred to th# leaeh aatar and (£) a eomtinaoa# treatment in ahieh
th# root# mould remain In th# eolation for a period of taelva hour#.
For each of the##, tao type# of treatment# concantratlom# of 1, 5, 80,
40 and 80 pim mere aaed.

For parpoae# of eomparlaon, tao Intermittent

aerie# mare in#laded a lth the concentration of 40 ppm for fifteen
minute# out of every hour and alao th# earn# period every tao hoar#.
When I t am# found that thee# tao aerie# prodaoed reault# almi X
ar to
thoae of the contlnuou# and short treatment# reapeotlvely, they acre
abandoned.
Early Investigator# have reported a periodicity In the mitotic
cycle In onion#.

Leal# (89) eonoladed that under condition# of normal

light and dark, there mere tmo aave# of mitotic activ ity , the maxima of
ahich came a t midnight and at noon.

Kelllcott (66) an# of the name

opinion but act the maxima a t 11 P.M. and 1 FÜi.
that the —
—

mod

Fklaaner (SI) reported

depended not upon the time of day bat rather

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IT
on tho tlm# of In itia tio n of notabolle a c tiv ity .

Thoaa eonolnolono

mar# roachod a fta r oboorvationo mara mad# of oampXaa takan at tmo hour
Intarralo.

Am a raault of th a lr Invaatl&atlom In mhloh root tlpa of

AUlmm mara aoUaetad ovary hoar and tho mmbora of division figuras
for a prodatanLinod araa mara ooontad, Solomon and Tj^ant (190) comaludad th at undar normal oondltlomo of day and night, thara mara no
daflnita mmvas of mltosas but rathar an hourly rhythmlclty of mitosis
in th is particular plant.

Thoir graphs shom maximal mitotic activity

at 9 A.M,, 8 A.M., 10 A.M. and 6 PJÉ.
In order to dataxmino tho ragolor mitotic pattern of tho root
tips troatod in th is aarlas of axporlmonts, thrao typos of controls
mara sat up.

(1) A consocutivs hour control for mhich root tip s mara

taken from an umtroatad onion a t tha ssmo tlmo that they mara coUoctod
from tho troatod bulbs.

(S) An individual root control mhich mas pra­

parad by taking 85 root tip s from tho ssmo onion a t tho ssmo hour and
preparing tho slides.

ITom th is group, tmclvo slides mars salsa tad a t

random for tho study.

(3) A sarias control for mhich a ssmpla mas

taken just bafora tho roots mara placed in tho traatmont solutions.
The data from a l l those controls arc included in Appendix I I .

Since

tho variation among tha roots taken from tha same onion a t tho samo
tlmo mas even gras to r than tha variation among those taken at consecu­
tive hours tho s ignif icanca of mitotic cycles may bo minimi nod.
Throughout tho an tira investigation tho onions mara kept under
normal conditions of day and night.

All axparlmonts mara conducted at

room tcmparatura, and, although th is fluctuated a l i t t l e , tho variation

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18

wmM not great enough to here had any appreeiahle effect upon the re aulte*

Daring the coarae of the reoearch the bolba «are atarted at

rariooa hoora of tha day and night, and the root tipa «are checked
before treatment for high mitotic aetirity #
After the preliminary, exploratory expérimenta, in «hich the
rariooa typea of effeeta «are obaerred, diaooaaed aad categoriaed, a
atamdard data aheet «aa prepared.

Table I I , page I t , ia preaented aa

a aample of the reaording method.
fa r each aet of treatmemta t«o root tip a «are taken every hoar
and placed on aeparate alidea.

In order to redoae the aobjectiv#

elamant, one elide fo r every hoar of eaeh treatment «aa a canted by
Dr. G. B. Wilaon, the d irecto r of th ia reaaaroh, and the doplioate
elide by the « r i t e r .
The prooedore fo r counting «aa atandardiaed bet«ean the t«o
obaervara.

Saoh elide «aa placed on the calibrated neohanlcal ataga,

the count «aa atarted a t a comata n t, predetermined point on the v e rti­
c a l axia and thia reading recorded om the data aheet.

Ccmaecutive high

pomered field a «are counted from edge to edge of the SS x 8S am cover
a lip and thoae to ta le are recorded in the table a aa "axia" to ta le .
The v e rtic a l axia «aa inareaaed by one unit each time and the count
continued u n til a t o t a l of approximately 100 divialon figurée^ «are
obtained from each elid e.

In the alidea lAere the 100 figorea «are

eahieved in the middle of the axia, the count «aa continued u n til the

ihere «aa one cxoeptlom to thla:
in order to equalise nimbera
in the caiae of the untreated bulb in «hich aeveral roota «ere col­
lected a t the aaam hour 200 diviaiom figorea for each elide «ere
counted.

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i

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§

so
r#m&lngm on th at axis bad b##n

In thoaa oaaaa whare, in

tha la ta r turars, tha dlTlaioa fl#para# «ara laaa noaaroua, i t aaa fraqnantly aaoaaaary to go bank orar tha allda and taka raadlnga a t tha
ona-half unit Intarvala.

In aararal oaaaa duplioata eounta aara aada

from tha aama allda to ohaok on tha oonaiatanoy of tha raau lta.

Sna-

aarlaa aara praparad of tha tao aata of raadlnga and thaaa ara praaantad in Appandiz I I .

Sinoa tao alldaa of tha root tlpa takan a t

hourly Intarvala in tao aarlaa of aaah traatmant aara prapar ad and
oountad and th la data la ta r oonbinad, tha hourly paroantagaa and araragaa in moat oaaaa aara baaad on to ta la ranging from 450 to SCO a alla.
I t ia raadlly raoognisad th a t, aaida from tha auhjaotlva alomant
mantiooad abora, tao faotora may influanoa tha oounta:

(1) tha araa-

naaa of tha diatrlbutioa of tha matarial and (S) ahathar or not tha
moat aotivaly dividing araaa of tha marl atom happan to hava fallan a i th­
in tha pradatazminad araa of tha count.

Boaavar, ainoa i t oan ba

aaavBMd th at a l l araaa of tha root t i p hava an aqual ohanoa of baing
avanly dlatributad mlthin tha pradatarmlnad araa of tha oount, i t ia
ballavad that th ia taohniqua may ba uaad to obtain an indioat ion of
tranda

ahawid ba Intarpratad aooordingly.

Sinoa, for tha moat

part, tha aavaral aarlaa uphald, within raaaonabla lim ita, undar tha
oonaiataney ta a t, i t aaa f a i t that tha influanoa of thaaa tao faotora
in diatorting tha piotnra oould ba minimi aad.

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OBSERVATIONS

Qomlltatlv# AapMta
Th# T#rlo#m type# of oytolosioal offoot# oa th# oaloa root t i p
■oriotao# of th# Aotl-dioa# t r o o f #at# ooa bo #ooa by or—laotloa of
tho plot##*

Tor parpooo# of ooaparlooa» Plato I oa# proporoA.

I t ooa-

tala# oorlo# of mltotlo flgoro# ohloh llla o tr a to Wmt 1# goaorally ooa•lAorod a# aoraally or##al#od flgoro#.

Tho la to ra l t Io o of a prophaoo

la P I. I , PI#. A, #hoo# tho ohroaoooBo# a# thoy roappoar o lth th alr tolophaolo polarity (PI# I , Pi## B) rotalaod alaoo tho prorloo# dlTlaloa# At
thl# #ta#o I t 1# th o o ^ t (156) th a t, a t tho klaotoohoro#, thoy aro a t l l l
la ooataot o lth tho aaoloar amabram# ovoa thoo#h, "oomalAorahlo ahlftla#
aay taka plaoo a# tho ohroaoooa# araa, ahloh fiU od tho aaoloar apaoo,
eeao to lo6#o thomaolvoo aoro or loo# a#ala#t tho a—bran#.*

Ia tho

polar proaotaphaao (PI. I , PI#. B), tho ooatraotloa of tho ohraaoaoao#
1# approxlaatoly ooaplotod aad th olr poaltloa# joot prior to tho aotaphaao plato allgamoat (PI. I , PI#. 0) aro ahooa.

MoraaJ aaophaao aopa-

ra tlo a , l . o . , klao too boro# f i r a t, oaa bo ao#a la P I. I , PI#. D, ohllo
la to r Ota### of anaphaao aoroaont appoar la tho tao foUoola# fl#oro#
(PI. I , Plga. X aad P).

Thl# anraaoat la ooaplotod la tho oarly tolo-

phaao of P I. I , PI#. G.

Tho roorgaalaatloa of th# d#a#ht#r aool#l,

la ltla t# d la PI. I , PI#. H, la ooaplotad la th# tao ##ta of adjaoaat
o#Ua la P I. I , FI#. I , ahoro tao latorphaa# aaolol oaa b# ao«e lyla#
to th# rig h t of th# lat# tolophaa# at###.

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as
Th# eytologleaJ. effect# of the Actl-dione treetmemt# mey be cab*
dirided Into too anjor eetecoriee:
intoreat boeanee

(1) thoee ohleh ere p rlaerlly of

of th e ir cytologleel end genetlcel p o te a tle lltle e end

(S) thoee which re fle c t verylng degree# of toxicity*
At the oeteet# I t meet bo pointed c e t, th a t the cytologlcel eberretlce# ehcwm in the pletee any be foond e# ccaperetlwely rero occwrr*
ence# In the entreeted a e to rle l (See deecrlptlcm o f the pletee) •

Fwrther-

acre, with the poeelble exception of thoee to x ic ity effect# which In re rlehly develop e fto r long period# of continnoe# trcetaemt, end with th#
higher eoneentretlcm# efto r treetaent fo r 15 alnetee, no pertleeler ebor*
rent flgorc con be eeld to be exeleelvely the re e e lt of e elnglo type of
treetaent, th e ir occwrrence being acre cloeely corrcleted with time them
with concentretlon*

These qeentltetlve eepeete of the problem w ill be

dlecueeed acre f e lly le te r e fte r e eervey of the eeverel type# of effect#
he# been aede.
There
tercet#

(1)

ere two mein type# of effect# ofp e rtleeler cytologlcel InThe orgenlsed flgere# which give the eppeereaee of hevlmg e

p ertlelly Inhibited eplndle end which, depending epon the degree of
eplndle euppreeeion, cem probebly re term to the noxael coédition In the
eeoceeding etege*

(S) The dleorgenlsed flgere# which re fle c t to te l In­

hibition of the eplndle

end which ordlnerily #e«m emeble to

return to the mormel pettom In th e t p ertleeler divielon*
The elngle type of orgsnlned flgere# ere thoee which hove been
celled "redeeticeel** grouping#*

A eerie# of cell# contelnlng "redeetloeel

grouping# ere eeecmbled la Pie te II*

In generel, these e oef Igeretlone

occur red meet frequently in the eerXy period e fte r treelment, l*e*, ebout

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23
four hours or loss*
Probably dopsmding upon tho s lto tio stogo of tho eoU a t tho tlmo
tho Aoti-diomo bosmmo offootlTo upon I t ,

thsro aro too possibiU tios of

tho origin of tho so "rodustlonal" groupings*

In tho f i r s t plaoo, i t i s

possiblo that thoso ooparationa, in sos&o oasos a t loast, may dorolop from
**nonal** figuras*

Ibr azsmplo* tho motuphaso# shoos In FI* I I , Figs* C

and D may hava dorslopod from an inhibition of tho middlo sootion of a
normal spindlo, and, lilcswiso, tho s p l i t onophaso in F I. I I , Fig* F may
hsTo foUosod a normally orgamissd notaphaso plato by a disruption of
tho usual by-polsrlty o f tho spindlo*

Qy tho ssmo tokan, tho s p lit in

PI* XI, Fig* H tolophaso may haro origimatod during or a fta r normal anaphaso moromont*
On tho othsr hand, i t s o i s oqually plausiblo th a t, dopomding upoa
suoh faotora as tho sin s of tho ooll, width of tho gap, and tho oriontations of tho groups of ohromosos&os, tha soparatioa may bo rotainod through
subsoqmsnt stagss of tho d irisio n and thus a dividod motaphaso plato
(PI* I I , Fig* C), uhioh may haro origimatod from a prophaso "rodustiomal"
grouping (PI* I I , Fig. A), nay, in tu rn , dorolop into s p lit snaphaso and
tolophaso figuros (PI. H , Fig* F, d and B)*
Thoso **roduotional'* groupings inrolrod oithor oqual (PI* I I A,
FTontispiooo A) or unaqual (PI* I I , Figs* B, 0 and D; FTontispiooo B)
ehromosono nsmbors*

In addition, notaphaso platoo wsro obsorrod in uhioh

tho groups of ohromooomos had oithor sim ilar (PI* I I , Figs* C and D) or
dissimilar oriontatioms (I^mtispiooo B).

This oriontation may bo a

factor in dotozmining uhothor or not tha separation would bo naintainod
in suooooding phasos of tho division*

I f tho two sootions of tho s p lit

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ZA
plat# ar# proparlj ariantad (PI* II* Pig#. C and D) i t is qui ta poasibla
tha gap bataaaa tha ahroswaoaa groups may ba alosad (partlaularily im
■ w llar s a ils ) as thay proaaad through tha msxt staga foimimg a normal
la ta amaphasa (PI, I , Pig. 7 ).

Im tha easas mtaara tha diridad grasps

hava diffaramt criamtations (Aromtispiaea Tig# B)* i t sa sms logioal to
assnsM th at tha saparatiom is mora apt to ba mmlmtslmsd im tha la ta r
stagss.

A scmparisoa of PI, II* Fig, F with P I, II* Figs, H and I mill

illu s tr a ta th is point,

Tha mida saparatiom batmaan tha iqppar groups im

tha f i r s t figura may ba maimtaimad so th a t tha following talophasa may
appaar similar to* or parhaps avam mora aosamtuatad than* that shoum im
P I, II* Fig, H,

Ihrthaxmora* dapandimg upon tha siaa of tha imtargroup

spasa* as mall as tha siza of tha sail* tha gap may aith ar r smsin saparatad or ba slosad during talophasa as shoum in P I, H* Tig, I ,

I f tha

l a t ta r os surs tha ra s u lt mill ba a raturm to tha ragular shrsmosos&a
nasbar.

I f tha gap parsists* homarar* thraa muslai mould ba fonsad* oma

with tha ragular and tmo with r ados ad sfaromosoma mmmhars,

Thaas mmslai

might prasant tha appaaranaa of a tria d sush as that illn stra ta d in
P I, VI* Fig, D.
Bridansa th at radusad muslai san ba fsonssd in a fam sasas foils# ing unaqual saparatisns mas found in tha ossasiomal ossurramsa of umbalans ad la ta talophasas (PI, V* Tig, IJ* of binuslaata s a ils with muslai of
unaqual aims (PI, VI* Fig, c) of trinmolaata s a i l s (PI, VI* Fig, D) and
tha vary raraly osourring radusad s a ils (PI, VI# Figa, T and F),

Tha

l a tt a r mara found im tha matarial takan too soon a fta r traatmant to hava
baan tha raaulta of tha Asti-diona influanoa and mora than likaly rsgraaant tmo naturally osourring "assidamts."

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ts
I a fli— i / t
1.

Th### "rvdAotlea#!" gpoAplAg# ##r# #h#rm#t#rl%#a by th#

pr##«nO# of #I#B#Bt# of OrgAAlAAtlOA.
£•

Th#y may bar# d«r#l#p#d from th# normal eomdltlon a t any atag#

In th# mltotl# oyola.
9*

Th# ##paratloon lnrolr#d both #qmal and naatimil mambor# of

ahromaoomaa*
4.

Tha ##paratad groupa had both alai la r and diaalmllar oriaata^

9*

Dopondlng upon auoh faotora aa also of th# ##11, th# orlaata-

tlo aa.

tio o , and th# uldth of th# gap batmaan ohromoaomo groupa tho aaparatloaa
may or may not bar# boon rotainod#
6,

Aa fa r aa th ia atudy la oonoarnad thoy gaaarally roaultod In

th# produotloa of nuolol mhloh had loat th o lr a b ility to dlrldo#
TUo typoa of dlaorganlaod figura# that appoar to bar# roaultod from
th# oomplot# Inhibition of th# aplndlo, namoly, th# "aogrogatloaal" group­
ing# and th# "aoattorad" arrangmaoata may b# rooognlnad#

l a thia roport

th* tarm "*a#gr#gatlonal" group la uaad to rafor to thoa# dlaor ganlaod
plat## In mhloh th# ohromoaomo# hava boon aoparatod Into tmo groupa#

In

oontraat to th# orgp"f aad "roduetlonal** grouping# #iloh ar# moat oharaotor
la tlo a lly found In th# oarly hour# after th# bogltining of traatmant, thoa#
mor# dlaorganlaod figura# mar# found more frequontly in th# lato r hour#
a fte r the i n i t i a l oxpoaur# to Aotl-dlono,
Plat# I I I oontalna a aarlaa of thoa# "aogrogatlonal" groupa#

In

P I. I l l , Pig. A and th# upper o o ll of Fig. B, oqual ma&bera of ohromoaomoa har# boon gpoupod.

Th# aoparation of th# oonatltuont ohrcmatlda

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B6
ia P I. I l l , Pis* 0 ha# baaa aaaplatad la PI. I H , Mg. D.

It

w ill ba obaarrad th at thl# moramamt d lffar# from th# normal tjp a , la
«hlah tha klaatochora# mora f i r a t (PI. I , Mg# D), ainoa hara tha ahrcmatida appaar to hara maraly fa llan ap art.

In tha abaanaa of an orlanta-

tion maahaniam. I t la highly probabla th a t th a j may ranaln mora or laaa
aloaa tagathar a lth tha raanlt th a t aaah grovp of ahrnnaaomaa may baacm#
inalmdad in a aiagla nnalana.

I f tha damghtar nmalai ara fovnad in thia

mannar, tha ra tn m to tha original atnrnmnanmi mambar w ill follow with
tha poaaibla ra an lt of ganatia aagragatiom withont a radnatiom in ahramo*
aoma mabar.
Aa nnaqnal (10-6) namariaal aaparation of ona of thaaa diaorgan^
iaad figaraa ia lUmatratad in PI. H I , Mg. D.

Tha poaaibla gamatiaal

aigniflaanaa of th ia w ill ba diaanaaad l a ta r .
In Sammary:
1.

Thaaa "aagragatioaal** gronpimga may ba aald to d iffe r from tha

"radnational" typa in th a ir laak of organisation, in baing Inaapabla of
ratnming to tha nrwmaT d iria io a pattam in tha naoct phaaa of d iriaian,
in baing fonnd mora fraqnantly whan a lomgar period of tlma haa al apaad
a f ta r tha

of traatmant and in affaoting aagr agatiom mlthorn*

radnatiom in ehromoaoma nombara.
Mamplaa of tha aaaond typa of diaorganisad fignraa, i . e . , thoaa
to idileh tha tarm "aeattarad* arrangamanta haa baan appliad ara ahoam in
tha fignraa of Plata TV and fTontiapiaea Piga. X and P .

Thay ara of par

tie u la r aignlfieanoa baeanaa, potantially, thay may land to tha polyploid
aonditlon.
from tha obaarratlona i t appaar# that th ia diaorganlnation may oeom
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S7

m t either propheee» metepheee or enephe— of n o n o l mitoel#.

y l,

Pig. At ehowe the reeo lte of e proaetepheee di«niptioB (Of. PI. I , Pig.B).
I t 1# quite poeelble thet theee prooetepheee dieorgeeisetione mmj here
developed from "effected, eœ-eegregeted" propheeee eueh ee thet ehoee
la PI# 7, Pig. 0.

Negleotlag the p o e e ib llitj meatloaed ebove, of groupe

of ohromoeoe&ee eegregetlag, there ere tuo p e e e ib llitie e of eubeequeat
developeeat of theee dieorgeaiaetloae.

Ia the f i r e t pleee, the prnmote-

pheeee m m j give rie e to "eoettered" metepheeee (PI. 17, Pige. B end C).
Aleo ohereoterietie of theee "eoettered” metepheeee ie the delej la the
e p littia g of the kiaetoohore ee ehoma ia P I. 17, Pig. D.

Homever, evemt-

uelly the e p llt mep ooeur, the ehrometide eppeer to f e l l epert (PI. 17,
Pig. r end PPoatiepieoe Pig# K) end, for e time, l i e p e relle l to one an­
other •

I f theee ehrometide ehould "d rift" epert the reeult oould be the

dieorgeaised eaepheeee euoh ee ehoma ia P I. 17, Pige. H end I end Proatiepieee P.

Xveatuellp e l l of theee ehrometide mmj be ineluded u ith la e

eiagle nmoleue end ia thie meaner the polyploid eoaditioa ie eetehliehed#
Thie hee been ehoma to here been the eeee e f te r treetmeat of oaioa root
t i p merieteme with eolehleiae (76).

The eeeoad poeeibilitp for the eub-

eequeat diepoeitioa of theee dieorgeaised pronetepheeee, eepeeielly of the
type ehoma la Plate 17, Pig. B, end eleo o f the "eoettered” metepheeee,
eaeh ee figured ia P I. 17, Pig. C, eometimee beoome "etieky" aad appear
to fuse together a t the ende reeu lt lag ia e "olamped" metepheee eueh ee
thet ehoaa ia PI. 7, Pig. P.

Xreatuelly the mhole group of ehromoeomee

eoatreete giving rie e to a re e titu tio a aueleue, mhloh, depending upoa
mhether or not the ehrometide eeperete may or may not beoome polyteae*
I t ie poeelble thet eometimee the dieorgeaiaetioa may take pleoe

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dnrlBc met&phm#e.
PI. IT# Tig. X.

Thlm &pp##re to Iulto ooevrrod in tho figure ohomm Im
In th is emeo# boemmoo of th e ir mppemremt mligment mt

m
m eqmmtoriml plmte# i t eeeme logleml to mmeimm thmt t ^

epirndXe imhibi-

tiome mmmt hmve oeourreA mfter the ehrcmoecmee bad ■mnetnrered to their
"mormml" metmphmae mligmnemt.

Xridenee of the obatrwstiom of the apimdle

meohani am m m j be found in the fmet thmt the ehronmtida mppamr to bare
fallen apart and laok the orientation of th e ir "noraml" aeparation (P1.I#
Tig. D).

Theee eella nay follow e ith e r of the pattern# outlined above

for diviaion figurea affected during prophaae.
Spindle inhibition nay alno oeeur in an early amaphaae c e ll auoh
aa ahoun in P I. I# Tig. D.

Aaaiailng th at ehronatida aeparated in a

normal manner# i.e .# kinetoeborea f i r at, and then the apimdle oeaaed to
function# the oriented chronoaome# may then " d rift" in a l l direotiona
and p ractically f i l l the cell# in a manner ahoun in PI. IT# Tig. O.
Thia aitnation may reau lt in the polyploid condition in the acme manner
aa th a t deacribed above for the unoricnted chroamtida that ariae from
the fa llin g apart of "Mattered" netaphaaea.

I t ia equally reaacnable#

homever# th at there may be a clmaping into more thna one group with
nuclei foxwMd f rom each gpoup and# ac a reault# bi aaa lea to (PI. TI#
Tig. C) tri-n a ale a te (Plate TI# Tig. D) or even amltinuclcate cella be
formed.

Nuclei formed in cuch a manner would be anauploid.

Theae "Mattered" netaphaaea occurred in the higheat frequency
after cxpoaurM to the higher concentrationa in both typea of treatmemta.
ytrthermora# there waa mo recovery from any of the treatment a in which
the "Mattered" netaphaaea apprnrrlmated 100)t of the total metaphaMC.

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s«
SlBo«a la th ia lavaatlgatlam, aaithar polyploid aor aaaaplold «alla
aara fouad that aoold hacra oeaurrad aa a ra a a lt of traatmaat. I t may
ba aaatBod th a t, uadar tha aomdltloaa proridad by tha laaah matar,
thara aara foraad ra a titu tlo a aa ala l, ahloh varlad In th alr ahromoaaata
awmbara and ahloh did mot divide a#ala#
la SwBwry;
1#

Thaaa dlaorgamlaad flyuraa ra fla o t aomplata lahlbltloa of

tha apladla.
8«

Tha dlaorsaalsatlon may oeoar a t prophaaa, mataphaaa or ana-

phaaa atagaa of mltoala*
3#

Sabaaqaant davalo]p»aat of thaaa dlaorganlaod figaraa probably

dapanda apon that ataga a t ahloh tha diaorganl zatloa oooar r ad#
4.

Lika tha "aagragatlonal** groapinga thaaa "aoattarad" figa

aara foamd mara fraquantly In tha la ta r abaarratlon parloda#
9«

Ravaralona to **raatlng** aondltlom aoourrlng wham oalla whlah

hava baan affaotad In tha aarly atagaa of mltoala raau lt In tha formation
of ra a tltu tlo n nuolol whloh may aomatimaa ba polytana*
Tha toadoity affaota of traatmamta may ba aub-dlvldad into thraa
aatagoriaat (a) thaaa whlah appaar ta a ffaat tha m
malaar mambrana, (b)
thoaa whlah aaam to affaat tha matrix raaultImg im a ohrnmaamo "atiokimaaa" and (a) datariaratlom of tha nuolaar oomatltaamta laocrlag only tha
apiral atruatura.

Xsamplaa of thaaa varloua taodo affaata ara aha#m In

Plata 7 and yypomtlaplaaa Tig. D.
Calla In whlah tha mmalaar mimbrana appaara to hava baaa affaatad
ara ahoam in PI# V, Plga# B, C, D aad E,

Whan affaatad la praphaaa, i t

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90
sppeered th m t tb m m m hrmnm may haw# baaa ratarAad la Ita a a tir ity ahlla
%ha ahramaamia# aoatiaaad to procaad through th a lr ragular anrphnlogloal
ohaagaa.

Latar tha aaabraaa appeared to here aoatraatoA dravlag tha aoa-

tanta of tha auolooa into a tig h t b a ll.
aa **naorotla* ia th ia atudy.

At th ia ataga i t ia rafarrad to

In PI. V, Pig. B tha ahromaaamaa aaam to

hara aahierad almoat th a ir mataphaaa appaaranaa whila a t i l l ooaapyimg
th a ir aarly grophaa# poaltioa (Of. PI. I , Pig. A).

In PI. V, Pig. 0 tha

ehromoaomaa hara raaahad th a ir mataphaaa length bat thay appaar to ba
"bald" imaida tha nualaar mambrana, and in addition, aoma hara loat th air
normal orientation aa ahoam in PI. I , Pig. B.

I t aoama quite raaaonabla

th at nualai of th ia typa may r a r a r t to intarphaaa in a mannar almi la r to
that ahomn in PI. T, Pig. D.

Brantually tha nualai of tha typaa ahaan in

PI. ▼
, Piga. B, C and D oontraat draming tha aomtamta of tha nuolaua to*
gather and tha "naarotlo" oomdition raaulta (PI. 7, Pig# X).

Thaaa

"naorotio" nualai aomatimaa may ariaa by rareralona of "aoattorad" mata*
phaaaa by may of tha "atiakinaaa" of ohromaaomaa aa ahomn in PI. V,
Pig. P.

Stagaa of thia rararaiom hara baaa obaarrad in thia atudy.
Tha aaaond tozie affaat mantionad abora appaara to inrolra a

liquifaation or diaaolution of tha matrix and tha ohromaaomaa daralop a
**atiekinaaa* manlfaatad by th air apparent fuaiona, f i r a t a t tha anda and
la ta r at pointa

th a ir length.

Thia typa of toocia affaat ia illun*

tratad in tha mataphaaa plmta of PI. 7, Pig. P, in tha anaphaaa of PI. 7,
Pig. G, and tha talophaaa of PI. 7, f ig . H.

Thara ia alao aoma of thia

"atiakinaaa" aridant in tha a p lit talophaaa of PI. I I , Pig. G.

In tha

laa of tha unequally aaparatad talophaaa groupa of PI. 7, Pig. I , i t

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31
###m# grob&bl# thmt th l# mltmmtlom oofold barm mrlmmm t r wm el* p ln c# r# •o ltla c from "«tiokiam##** la m dleeopsmalsed mamphm##*

I t 1# peaalbl#

thmt #uoh m flgmr# might rmmolt in m hl-mwlsmtm ##11 #uoh ## thmt shomm
la P I. VI, Pig. C.
Th# dmtmrlormtloa of th# aoalmmr ooamtltumat# 1# mth ird typ# of
toad# offoot ob##XT#d la th# eonro# of thl# #todj.

Th# #plrml#d ohopom-

oamomtm olom# mppomr# to b# romlotmat to thl#, aad ma m ra a a lt, la th#
##11# mff#ot#d la thl# mannar oftaa th# aplrml atr aotar# booomam «ridant.
Pfaatlaplao# Pig. D mad PI. VI, Pig. O ooatmln tolophma# flgorma la ahloh
th# aplrml# or# «ridant.

Th# latarphm### mhoam la PI. V, Pig. A giro ma

Indlomtlom of ahmt hmppaa# ahaa toodo ooadltlon# mmnlfaat thamaalra# mt
thl# ataga.

Har# I t mppamr# thmt th# eonatltaant# of th# aaalaa# hmr#

datarlormtad a a t l l only th# amoollad ohromomammtm romaiaad g lrlag th#
naolaa# a "thraady" mppamraaa# mot aallk# thmt of th# sygotam# aaalaa#.
Th### latarphm### mr# qalt# typloml of th# Imtar hoar# aftar t r aatmant
a lth htghar oonoaatrmtlon#.
Ia Ammmry:

Th# foUowlag aara ooaaldarad m# Indloatloa# of toodo

ooadltlon#.
1.

Imtarfarano# with th# aaolamr mambrana aad aramtamlly emaalag

a oontrmotlOB of I t .
£•

Chromo#om# "atlokln###" manlfaatad by faaloa#, by atrmada ax*

from ohoromoaom# group# and by unaqual aaparation#.
5.

Dlaintagratlon of th# eontaat# of tha mmolau# and/or th#

ohromomcm# matrlT la dividing oalla.

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58
A group of MioooUunoooo oboorvutlou#, moot of uhioh or# oouporutlToljr iafroqu#nt, eompoo# Plato VI.

With the ezoeptioa of the

blBueleate (PI. VI. Tig. C) and the triaueleate (PI. VI. Pig. D). the
figure# la th is plate map be aald to have been observed onlp rarelp.
Theee figures, as u e ll as the reduced c e lls (PI. VI. Figs. g and F)
have been dlecueeed.
The polpplold esephase (PI. VI. Fig. A) occur r ed in a root tip
taken three hours a fte r exposure to 1 pjm fo r IS minutes.

Since su ffi­

cient time had not elapsed a f te r treatment, th is oould not be attributed
to an e ffec t cf Actl-dione.

I t Is sig n ificant in th is studp as an indi­

cation th a t, cccasionallp. polpplold c e lls are produced in the "ncsmal"
course of events.
Another occasional observation

uas ehrcmoeoms fregsentatlon

(PI. VI. Fig. B and Frontispiece Fig. C).

Since these fregpcenfed chrom­

osomes appeared so rarelp In the slid e s. I t seems most llkelp that thep.
lik e the single polpplcid c e l l mentioned above, also msp have been
"natural" accidents.
A destruction of the la te ans phase or ear Ip telophase polar Itp and
consequent separation of the chromosomes map have been responsible fcr
the peculiar telophase shomn in PI. VI, Fig. G.

I t might be of interest

to note th a t th is figure contains an indication of toxleltp In the revela­
tion of the co il structure.
The c e ll In PI. VI. Fig. H map be classified as a sort of "tran­
sitio n al" metaphase.

I t appeared in a root t i p mhich mas taken after

three hours exposure to SO ppa of Acti-dione and appears to re flec t a

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99
traiMitiom between a s p lit metaphae# anA a aomplataly Alaoreanlsaâ
oaa.

Tha delay in tha sp littin g of tha kinatoehoras saomad to aa-

oompany tha obstmetlon of tha spindle and th is delay ia evident in
tha figura»

I t seams reasonable, therefore, th at there may have bean

a p a rtia l inhibition of the spindle a t f i r s t and tha s p lit mataphaaa
developed, th at th is mas foUomad by the suppression of the astiv ity
of the rsmaindar of the spindle.

As tha s p l it metaphase started to

f a l l apart the root tip mas removed and fixed and thus the s e l l mas
caught in a staga tran sitio n al batmaan an organised s p lit mataphaaa
and a disorganised arrangement of aith ar the "sagragatiomal" or
"saattarad" typa.
The laak of orientation of tha ehromatids ia an indioation of
tha

disorganisation in the o all im PI. VI, Pig. I#

Beoansa of tha

olmmpiag in tha eenter, i t mas impossible to eount tha ehromatids
smasesd into the group, but there appeared to ba a radusad awbsr.
On th is basis i t is intarpratad as an unipolar anaphase of a radusad
o a ll.

However, baoausa i t appeared in a root t i p ta ken only tmo hours

a fte r exposure to 80 p%m of Aeti-^ione, only the disorganisation, and
not the raduotiott, may ba oonsidarad as a ra su lt of tha antibiotie
traatmant.
In Timmisry;
1.

Polyploid s e l l s , fragaantatlorn of ehromoaomaa, radusad

c a lls and tran sitio nal stages batmaan tha several oatagorlas desoribad mere vary rare oaourransas la tha observât ions.

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34
A#p#et#
Th# r##*lt# obt#lii#4 ia th# q##atlt#tlv# ##p##t# of thl# otaây
#r# pr###Bt#d ia th# toblo# of AppoaAix I I .

fo r purpo### of pr###at#-

tio a th# data w ill b# growp#d iato two oatagori##:

(1) Thoa# which

r#fl#ot th# goacral affoct# of th# Acti«dioa# on th# mitotic cycle aad
in#lad# a coacidoratioa of th# to ta l acmbcrc of divlcioa figure# a#
wall a# th# comparatiT# fraquaacic# of th# iadlvidual #tag##; (I) Tho#c
which indicate th# more cpccific effect# of th# traatmaat a# maalfaatcd
by th# ooccrraaca# of each of th# type# of abarratioa# di#cu###d ia th#
praccdiag ccctioa.

Th# ccmbiaad raault# of th# f i r a t five hour# a# well

a# th# hourly variatiom# of th# affect# of th# Aoti-dicm# treatmeat#
w ill be compared,

la aeveral eerie# of traatmaata, (particularly th#

ehort) a f ta r a period of five hour#, there war# iaoufficiaat diviaioa
figure#^ fouad ia the entire elide to obtain th# minimum count of 100
(Appendix I I , Table# 1 through 10).

Tharafora, a compariaoa of th#

combined reçoit# of a l l treatmeat# may only be mad# oa tha baai# of tha
f i r a t five hour# a fta r axpooure.

Accordingly, to equalise th# data for

ouch a comparicom, ocparata TOhla# I I I , IT aad T have bean prepared.
A. Oaaaral I f f act# of Acti-dioae
Tha average aimibar# of diviaioa figure# per azia* obtained from
tha combined data of tha f i r e t five hour# ara ahowi ia Table I I I .

Vbaa

^ Tha reader i# referred to PI. I , Piga. A through H to obaerve
the range within which tha call# were coaaitered a# being ia th#
atata of diviaioa.
* "Axia" refer# to tha eoaaecutive aerie# of high pomered field#
takam from edge to edge of tha cover a lip .

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55
th#

tr c m a l l

maWrlal mr# eaeeilnad, (Appaadix Tmbl##

SC through sec), i t w ill bo oboorrod th a t 16114 dirioioa figura# war#
found cm 9S6 axaa giving tha araraga of 17.5 dividing aalla par azla*
Thia aoBparaa f avor ably with tha naan (SO.S) of tha wmtraatad root#
takan a t eonnaoutiva hours for tha f i r a t f iva hours.
TABLE I I I
AVERAGE HOM
BER 07 DIVISION 7I0DRBS PER AXIS OBTAINED
IRON IBB CGMBINBD DATA OF FIRST FIVE BOORS

Oonoaatratloa

Tÿpa Traatnant

1 m

Short
Oontinmous

5 PRC

Short
Oantinuous

so PRl

Short
Oantinuous

40 PPM

Short
Continuous

80 PPM

Short
Continuous

Ohtraatad (Oooaaautiva hours)
All Dhtraatad Material

Total Count

Anas

Avg.

18.6
15.S
10.8
14.1
11.6
S4.5

1774
SS15
S544
SlSl

19S
191
SS5
18S
195
100
S46
155
187
179

S4S1

119

S0.5

16158

9S6

17.5

S4SS
S5S4
S4S1
S574
SS69
S446

7JI
14.5
18.5
11.8

Tha data of Tabla I I I ara graphioally prasantad in Tazt Fig. 1.

Trcm th is histogram i t is avidant th a t, on tha basis of tha avaraga
ntabara of division figuras par axis obtainad from tha aonbinad data of
tha f i r s t fiva hours, tha re su lts of tha two types of traalmant nay bo
said to have beam similar a t aithar and of tha aonaantration sarias with
a divarganaa oaaurr ing in tha aanta r and tha graatast diffaramaa be twain

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5#
%h#

of %h# too type# of t r e e f e o t eppeerlog e t the tO ppe

eoneeotretion.
Before e eooelderetloo of the hourly Terietloo la the everege
mmihere of dlTlelcn figeree per exie eoald be umdertekem. I t eee aeeeeeery
to obteln iaforoetloa eoaeeming the Tarletloa emoog eererel root#
from the eeme oolom e t the eeme hour.

Aeoordlagly, the ladlTlduel root

eomtrel me# prepered (8#e deeerlptioe of method, pege 19).

The dete

token from the root mhleh heppened to here been ehoeea f i r et eee erbi*
tr e r i ly need e# e beee end the derletlom# of other ererege# from i t mere
eeleuleted (Appendix I I , Thble SB).
(Text

When theee derietiom# ere grephed

f i g . 8), one een reedily eee tb e t not only 1# there e eomeldereble

Terietion —

# the IB root# in the meen nm&ber of dieieiom figure# per

exie, but eleo there le e wide renge from lomeet to hlgheet point
(+18.8 to ->7.S).

Attention ie celled to the fe e t thet the hourly verle-

tione, sheun In the graphe (Text Flge. B through 9, 9 through IS), ere
euperlmpoeed upon en underlying pettem of wide Terietion from root to
ro o t end thi# feet meet be kept in mind when Interpretetion# of the dete
ere being mode.
In order to etudy the Terietion In the ererege nmher of dirleion
figure# per exie, the meen of the mere hour ( i . e . , juet before the tre a t­
ment wee epplied) wee need e# e be ee end the derletion# from thie point
were oeleuleted (Appendix I I , Table# B9 through SB).

When the derietion#

from the mere hour ererege of the untree ted meter le i ere plotted in their
hourly eequenee (luxt Fig. B), owe mey obeerre th et the range of fluetuetic n wee lee# (+10.1 to -9.0) then i t bed been fo r the root tip# ooUeete^
from the eeme onion e t the eome hour (+18.8 to —
9.S).

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

In addition, the

87
hoorlx vmrlmtlon Im mo#t e##em «m# 1### than that oaenrrlnc nooc roots
taken a t one tla e .

Although both of these graphs were aade on the basis

of approximately the seme atsnbsrs of c e lls , each point in Text rig .

e

Is

based ott readings taken from a single root (aee method, footnote, page
IB), mhile the data for Text ? lg . %were accumulated from four roots of
two d ifferent onlcms*

Increasing the mwaber of roots and of celons was

the method used to lower some of the radical peeks th at generally occur
as a re su lt of individual variation.
Text Figs. S through 7 end 6 through 19 have been constructed in
a manner similar to that described fo r Text Fig. 8.

In those cases where

the division figures were insufficient for the counts to be made, the
condition Is indicated by a dotted line ending at a point below the
abscissa equal to the average nmber of divisions found before the In­
itia tio n of treatment.
After the short (IS minute) treaimmnt with the 1 ppm solution,
the re su lts produced (Text Fig. 3 ) , a t f i r s t glance, appear to hare been
scarcely d ifferent from thoee obtained from the untreated root tip s (Tmoct
Fig. 8 ).

Bowever, the peak that developed a fte r eight hours la this

series Is rather interesting.
normal range.

Most of theee divisions were within the

This sodden spurt of normal division was rather striking

and one that occurred in both roots of both series (Appendix I I , Table
16).

Although th is difference was not greater in range than that occurr­

ing bet ween two separate roots In the Individual root control series
(Text Fig. B), i t was larger than that observed between any two untreated
roots taken a t consecutive hours (Text Fig. 8).
Aside from the re su lts obtained from exposure to 1 ppm for 18

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97
hourly Torlatlon la moot emooa mm# 1### thma that oeemrrlmg —«mg root#
tmkon a t oa# tlmo.

Althoagh both of th##a graph# mar# #md# on th# ba#i#

of approzlmatoly th# #ob# mw*b#r# of call#, aaoh point la T#zt Mg. 8 la
ba##d cm raadlag# takaa from a alagl# root (oo# mothod, footaoto, paga
18), mhila tha data for Tazt Mg. 9 mara aooimiulatad from fonr root a of
tmo dlffaraat oaloaa.

laoraaalag tha aambar of root# and of oaloa# maa

tha mathod uaad to lomar aom# of tha rad leal paaba that gaaarally oaamr
aa a raault of ladlTldmal variation.
Tazt Flga. 5 through 7 and 8 through 19 have baan oonatruotad in
a manner almilar to that daaerlbad for Tazt Mg. 9.

In thoaa oaaaa mhar#

tha dlTlaioB figure# mara Inaaffielant for tha eounta to be made, tha
aondltlon 1# indicated by a dotted line andlag a t a point balom tha
abaolaaa equal to tha avaraga mmmbor of dlvlalom# found before tha In­
itia tio n of treatment.
After tha ahort (15 minute) treatment mlth tha 1 ppm aolutlca,
tha raault# produced (Tazt Mg. 3). a t f l r a t glance, appear to have baan
acarcaly different from thoaa obtained from the untreated root tip# (Tazt
Mg. 9).

Bomavar, tha peak that developed a fte r eight hour# In thl#

aarla# 1# rether Intaraetlng.
normal range.

Moat of theee dlvlclon# mara mlthin tha

Thla auddan apu rt of normal division mas rather striking

and one that occurred in both root# of both aarla# (Appendix I I , Table
16).

Although thl# dlffarenoa me# not greater in range than that occurr­

ing batmaan tmo aaparata root# In tha individual root acntrel aarla#
(Tazt Mg. 8), I t mas larger than that observed batmaan any tmo untreated
root# takan at conaacutiva hour# (Tazt Mg. 9).
Aside from tha result# obtainad from azpoaure to 1 pjm for 19

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38
miaut#m «hieh r##ambled thOM obtalaad in control

and, d##pit#

th# two nnd#rlying varying pattam# obaarvad In nntraatad m atarlal, tha
two type# of traatnant may ba aaparatad on tha baaia of tha trand# ob­
aarvad In tha hourly ohangas la tha avaraga mmbara of dlvlalom flguraa
par azla*

Following tha ahort traatnant a ganaral trand toaard a radua-

tloo in tha avaraga mmbara of dlvlaion flguraa naa avidant (Tazt Flga*
4 th ro u ^ 7)*

On tha othar hand, undar tha oondltloma of oontlnuoua

axpoaura to Aotl«dloma, two d la tln o t pattam a wara nanlfaatad (Tazt Flga*
9 through 13) t

Tha 1 p%m and 8 ppm eomoantrationa (Tazt Flga* 18 and 16)

diaplayad tha aana trand a aa gomarally appaarad following tha short tr a a tnanta*

In tha 80 ppa and 80 ppm oomoantrationa of tha oontlnuoua aarlaa,
■a

tha ganaral tandanoy was an inoraaaa in tha nwsbara of dlvlalom flguraa
i n itia lly and la t a r a fluotuatloa around tha avaraga masbar th at wara
prasant a t tha baglnulng of traatnant*

Tha 80 ppm aarlaa foUowad thla

ganaral p attam during tha f i r s t olght hours a fta r whieh tima tha dlvlalom
flguraa diaappaarad*
Sinoa tha aama oonoaatratlona wara uaad In both tha ahort and oon­
tlnuoua traa^&anta, l l t t l a dlffaranoas might ba axpootad to hmva ooourrad
a t tha and of ona hour*

Whan tha graphs of tha aama ooneantratloma in

tha two typaa of traatmanta ara oomparad, ooa finds th is to ba trua In
a l l oaaaa wzoaptlng tha 8 ppm aarlaa Wiara tha two typaa of applloatloma
produoad opposing raaulta*
Whan oomparlaona of trand a raault Ing from tha two typaa of tra at­
manta ara mada on tha basis of tha hcurly variation during tha f l r a t flva
hours a f ta r azpoaura to tha d lffa ra n t oomoantrationa of Aatl-dloaa, d lf f aranoaa In tandanolaa baooma appmraat*

In tha oasa of tha 1 P ^

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aarlaa.

S9
th# mmmm sanaral tr#ad# app#mr#4 fo r th# T i n t thr## hour# hut u t th#
fourth hour th#y dlr#re#d mud th# urorug# uimb#r# of dlTloio# flgur##
inor####4 mft#r th# short tr##tm#nt uhil# thoy sturtod # mmrh#d d##liu#
in th# ooutlnuouoly «xpoo#d ##rl##«
of th is •itu ntio n uu# found,

I s th# 5

tr##ts#mt#$ th# r#r#ro#

Oppooinc trond# #pp##r#d during th# f i r s t

tuo hours snd, s f t s r th# third hour, th# ##■■ psttorm us# folXousd in
#s#h of th# tr#stm#nts.

With 2 0 p%m diff#r#n### in th# rs s u lts obtsinsd

from th# tuo mothod# of sppXiestion sppssrod,

Thllouing th# short t r s s t -

m#nt, th# gr#st##t nmbors of division fig u rss usr# rssshod s t th# #nd
of th# f i r s t hours th is insrsss# uss foXlousd hjr s stssd j dsorssss u n til,
St th# snd of th# f i f t h hour, thsr# u#r# in su ffio isn t fig u rss to obtain
th# minimum oount of 100 dividing esU s on th# o n tirs slid##

Dhdsr th#

ooadition# of oomtinuou# srposurs, mi ths sontrsry, th# svsrag# mumbors
of division fig u rs s stssd ily ross fo r tuo hours snd than f o il nor# grsdu slly , but did mot d##lin# nuoh bslou th# svsrsg# of th# nsro hour#

Whsn

th# 40 ppm oenosntrstion us# ussd, i t u iU bs obssrvod th s t thoss g#n#rsl
t#nd#noi#s u#rs osrrisd fu rth o r, i# # ., thsr# uss s stssdy dsolins in ths
nombsrs of div isio n figurss s f t s r ths short trsstm snt u h ils , in thoss
roots coatinuously szpossd, ths nssn nnmbsr of dividing esU s, s f t s r sn
i n i t i a l inorssss in th# f i r s t tuo hour#, rsturnsd to spproximstsly th#
asms s# i t had boon s t th# bsginning of trsstmsnt#

Tbs trsnds obssrrsd

during th# f i r s t fiv# hours following origin al sxposur# to th# 00 p ^
solution usr# somsuhst similar in both ssrio s of tro s tuants, ths major
differsno# bsing in ths tim# ths pssk of division figuras us# rssohsd#
I t w ill b# obssrvod to hsv# boon s tts in sd on# hour s s r lis r s f ts r ths

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40
•hort t r — tm m n t tbm n i t

im tb# roots oootiimoaslx la ooatmot with

th# ##m# eomoontro t lorn o t Aotl«dion#*

*^am thorn# rooulto i t « i l l bo oboorrod that, on tho booio of ororog# ntmboro of division figurss por s%is, tho f i r s t f ir # hours msy bo
oonsidorod to hsvo boon tho e r l t i e s l porlod fo r tho oomporstiro obsorvstlon of tho difforsnoos in offoots of tho sovoml ooaoontrstioas of oooh
typo of trosWont*

In ni— s r y ;
T o r untrostod mstorisl, thoro uss s eonsidorsblo rsrlstio n

1.

in tho crorsgo numbors of division figurss per axis in tho individual
root control as uoU as in tho eonsoeutivo hour control sorios.
£•

Dospit# tho underlying pattern of variation in tho controls,

the treated material shooed definite trends im tho moans of tho nwbors
of dividing c o lls.
9.

follouing tho short troatment with 1 ppm, for a ll praotioal

purposes, tho results obtained uero not nnlilro tho controls.
4.

Thoro uas a general doelino in tho numbers of dividing eell

follouing exposure to a l l eoaeentrations of tho Aeti-^iono for 19 minutes
as u e ll as during tho period of continuous comtaot with 1 ppm and 5 ppm
solutions.
9.

Thsre uas an in i t i a l rise uhich uas foUouod by a decline and

a leveling off at approzimatoly tho starting point under conditions of
continuous treatswnt to tho SO ppm, 40 p]p& and 80 ppc comoontratioms.
In tho la te r hours of tho 80 ppm sorios division figures disappoarod.
6.

Tho f i r s t five hours a fte r tho in itiatio n of tree Went uas

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41
th# mo#t e rltio a l period for th# ob##rv#tlom of th# #ff#et# of th#
##ver#l tr##tm#mt# on th# froqnonoy of dirlslon flgor##.
Another npprooeh to th# etody of th# goneml effect# of th#
A#ti-4ion# trcntnont# m#y h# found In compnrleone of th# fr#qo#noi#e of
th# Individuel etegee*

for thle phee# of th# etudy th# pcrccntegee of

th# to te l nmmbcr of dlvlelon figure# «ore oeleuleted for propheee, metephee# end poet-metephee# etegee.

Thee# peroentegee ere preeented In

Appendix I I , Thhlee 1 through 15#
In th# "SoHBery of e l l untreeted meter lei" Included In th# etudy
(Appendix IX, Teble 19C) I t mey he ohecrrcd thet propheee compoecd 49)(,
metepheee t4#8j(, end poet -metepheee S4#8J( of th# 16,114 dividing celle
counted In th# untreeted meterlei#

In order to obteln the proportlcmet#

hourly verletlone In theee etegee, e l l pcrccntegee mere converted to e
coemwm bee# of 90/% (Appendix I I , Teblee ID through lOD, Teble ISD end
Teblee UB, ISB end 14B).

Thee# hourly couverelon figure# mere plotted

end the grephe d l l be found In Text flge, 14 through t7#
Text fig 14 ehoue the proportlonete hourly verletlone obeerved In
the untreeted bulbe#

In thl# greph I t d l l be eeem thet cech of the

etegee fluctue ted eround the bee# of SO#

The eeme nee true In the ceee

of the untreeted root# uhlch mere tehen from the eee# onion et the eeme
time (Text fig . 25).
When the proportlonete hourly verletlone In theee etegee ere com­
pered, dlfferenoee d l l be obeerved, not only between the two type# of
treatment, but eleo
tlon.

the coneentre t lone of each method of eppllce-

The goncrel reeult of the ehort treatment wee an Inoreaee In the

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48
propha###* tout th# pattern of tha iaerea## diff#ra eoeeehat —

g the

ooae#ntrations ns#d«
Aa a re s u lt of exposure to 1 ppm sad to 5 pp& for 19 minutes
(Text Pigs. 19 and 14) the very marhod inerease mhloh oeonrred during
the f i r s t two hours a fte r the treatmiant was In itia te d , was followed by
a dee line In both eases.

At the fourth hour, however, these two eon*

centrations seem to have prodused different effects%

In the 9 ppm

solution series the prophase proportion continued to decrease u n til the
divisions ceased a t tha end of the seventh hour, while in tha 1 ppi
material there was a second r is e In prophase frequency which began at
the f i f t h hour and was more or less retained u n til the end of the eleventh
hour.

The general r is e in the prophasec assiamed a different pattern In

the rmmalnlng three concentrations (Text Pigs. IT through 19).

In these

cases, the prophase Increase was a more gradual one, and the highest peak
In each, which Is practically the same for each of these higgler someam*
tre tlo a s In the la te r hours, was achieved a t the la s t hour. I . e . , just
before the divisions ceased en tirely .
I t Is to be expected that the metaphase and poct-metaphase patt*
ems would fluctuate with the prophases.

I t i s interesting to notice,

however, th a t, depending upon the comeentratIons used In these short
treatments, there was a divergence la the effects upon the frequencies
of these stages.

The 1 p]^ and 9 ppm solutions of Aotl*dlcme resulted

in an i n i t i a l decline In metaphases and assumed different patterns In
the la te r hours (Text Pigs. 19 and 16).

In the 1 ppm treatment, after

the i n i t i a l drop, the metaphase fluctuations were comparatively slight
and there was a tendency for a leveling off about 10^ below the

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45
p#re#ntmg# pr###nt prior to th# treatment.

In the 5 ppn oooeentretlom,

homever, a fte r the I n i t i a l drop» there oa# a rather steady rise in meta­
phases u n til the f i f t h hour and, following a sharp dee line (21.9^ in one
hour), there uas an equally aeute inerease (22.8# in one hour).
The metaphase ploture began to ehange someshat in the 20 ppm
series (Text Pig. 17).

In th is ease, i t may be said that the general

pattern «diieh resulted from the 5 ppm treatment uas foUoued but the
changes uere more gradual ones*

I n itia lly there uas an almost negligible

rise (4.1#) in the frequsney and the decline, uhieh took am hour langer,
was sim ilarily followed by a second increase.
A radical change in the effects of the ahort treatment with Actidione on the metaphase frequencies occurred when a 40 ppm solution was
used (Text Pig. 18).

There was a very marked increase in the frequency

of th is stage (from 90.0# to 96#) during the f i r s t bso hours following
the in itia tio n of treahaent and th is uas followed by an even sharper drop
(from 96.0# to 16.4#) in the second tuo hours.

I t w ill be observed that

these trends of metaphase frequencies is exactly opposite to those which
developed during exposures to lower concentrations for the short interval
(Text Pigs. 16 through 18).

This early dominance of metaphases, a l t hough

not continued, i s considered a rather important characteristic of the
resu lts of th is treatment.
After the short treals&ent with the 80 ppm the metaphases followed
the same general trends but the hourly changes, although s t i l l somewhat
marked, were more gradual ones (Text Pig. 19)#
In the two lowest concentrations of the short treatment i t is of
in terest to observe th at the post-metaphases followed the seme general

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fr#q%wnoy oc*r## m th at of tho motaphoooo.

In othor aordm, a# the pro*

phases Inoreaead o r deoreaaed, both of theee stages, for the most pert,
f e l l and rose acre or less together.

Sine# in the nntreated roots these

too phases oere found In almost equal pereontages, i t appears that both
of these la te r stages of the mitotic cycle oere almost equsilly sharing
the effects of the Aoti-dion# in these looer ccnsentrations.

After the

short treatment o ith the 20 ppm solution, however, th is oas not the case
and the metaphase and post-wetaphase frequencies began to diverge oith a
sharp ris e in one being aecosipanied by a correspondingly sharp decline in
the other.

This trend, ohich f i r s t appeared in the 20 ppm series, and oas

even mere accentuated in the root tip s ohich oere treated o ith the 40 ppm
solution for the short period oas also observed following the 80 ppm
treatment.

After the f i r s t too hours in 20 ppm of Acti«dione, a stsady

decline in the post-metaphase frequency oas observed, but in the 40 ppa
and 80 ppm groups during th is same period, sharp increases of this stage
occurred.
In stommry, th is study of the proport ionate hourly variation of
the individual stages following the short treatment oith Acti-dicne
revealed the following:
1.

After exposure to the 1 ppm solution, differences in the

relative frequencies of the individual stages began to appear from five
to six hours a fte r in itia tio n of treatment.
2.

The general effect of tha short exposure oas a ris e in the

relative frequency of the prophaaes.
3.

n iis prophase rise mms rather acute in the early hours foliota

ing exposure to the looer concentrations of the antibiotic and more

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45
gradual la th# hl^i#r oa#a«

Mataphaa# fr^quaaej fluotuatad aonaldarahly aa a r#sult a t

4.

th# 15 mlaut# azpoaura to traataaat aolutlona and haaama th# dominant
faatura In th# aarly hour# following th# ahort traatnant with th# 40 ppa
aolution*
5*

Poat-#Mtaphaa#a tandad to follow tha mataphaaa pattam follow­

ing azpoaura to tha lowar (1 ppm and 5 ppw) aanoantratlona but dlvargad
aonaldarahly a fta r hlghar (SO ppm, 40 ppm, and 80 ppm) aoneantrationa and
dcmlaat#d th# aaan# batwaan th# third and f i f t h hour a fta r aontaat with
th# 60 ppm aolutiom for 15 minutaa.
Tazt flga* SO through S4 ahow tha proportlomata hourly variation
in th# paroantag# fraquaaaiaa of tha IndiTldual atagaa in a a ta ria l aontiauoualy traatad with tha aavaral aomeantratlona of Aati-dlona*

Whan

this oat of graph# la oomparad with thoaa of tha ahort traatmant (Tazt
figs* 15 through 19), tha 1 ppm diagram of th la group Imaadlatalf stand#
out aa baiag mora almilar In Ita ganaral trand# to thoaa of tha Inwaat
thra# eon#antratIona of tha ahort azpoaura aarlaa*

In thla oaaa (Tazt

Fig* 20) th# pradomlnanc# of prophaa## and I ta mor# or laaa ataady,
gradual rla# w ill ba notlead to ba quit# similar to th at obaarvad In th#
20 pjm ahort traata&ant graph (Tazt Fig. 17)*

Th# mmtaphaa# and poat-

nataphaaa llnaa im thaa# two graph# also hav# raaamblamoaa In thalr gan­
aral oouraaa*

I t will ba obaarvad th a t, although thay follow th# aama

pattam . In th# oaa# of th# aoatlnuoua traatmant tha ohangaa wara aora
gradual om&aa.

Thla tandanay toward mora gradual ohanga whloh was aanl-

faatad f i r s t In th# highaat oon#antrat lea of th# ahort traatmant la th#

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46
mo#t predmleatliie f##tnr# of the eontimKrae tree^Mnt series efter the
f i r s t hoar (Text fig* 19)*
The 9 ppm eontianooe treetmeat (Text f ig . 21) eppeers to hscre
not only some effeote sim ilar to those prodoeed by the higher ooaseatratiOBs a fte r 15 aiantes exposure, bat also some re su lts reesmbliag those
observed ia eomtiaooasly treated material, and eould, perhaps, be oomsidered intezmediate la thla respect*

The effect a simulated those resulting

from the short treatment ia the following features:

(a) the disappearance

of divisions within twelve hours and (b) during the f i r s t tlre e hours, the
gradual rise in prophases, the sharp decrease in mete phases and the almost
equally sharp decline in poetisetaphases are features uhich were also
pointed out in the re su lts of the exposure to the 40 ppm solution for 10
minutes*

On the other hand, the effects th at d iffe r from those of the

short treatment are:

(a) the more gradual changes, (b) the dominance of

metaphases in the la s t three hours when divisions were present, and (a)
a fte r th e ir i n i t i a l drop, the slig h t variation in the relative frequencies
of postHsetaphases*

In other words, of th is particular treatment i t might

be said that during the f i r s t two hours the re su lts showed tendencies that
appeared eimilar to those of the higher concentrations of short treatment
and, in the la te r hours, the effects more nearly reesnbled those obtained
In the continuously treated group*
The sim ila ritie s in the trends of each stage in the graphs of the
20 ppm, 40 ppm and 80 ppm series of the continuous exposure to Asti*
dicne (Text Tigs. 22 th ro u ^ 24) are most strik in g.

I t may be of interest

to observe th a t, a fte r the f i r s t hour, there was l i t t l e hourly change in
the frequencies of any stage and that the tendenciee established at the

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47
•nd of tho f i r s t two hears wsrs mors or loss rotsiasd throughout tho
rsmsimlng period of tho oboorrstlom.

Tho out stood leg footuros of those

higher eonoontrotioms of oontinnous trootmont more tho domimsnso of tho
Botsphosos, tho rsd io sl roduotiom im tho post hsstophssos sad tho more or
loss stohlo proportion of prophoso figures.
ppm sorios of th is group (Text Fig. £4):

Chongss sppssrod in tho 80

(s) Tho divisioms dissppesrod

s t ths snd of sight hours; (h) Tho motsphsso r is e mss mot ss grost ss i t
hsd boon during ths oompsrsblo period in ths 40 pps solutiom; (o) Thoro
mss s grostor goaorsl drop in tho prophssos.
When ths rs s u lts obtsinsd in tbs ssriss using tho highest somosmtrstioms of ths oontinuously szpossd group sro oompsrod mith ths ssmo
oomeontrstion of ths short trsstmsnt sot, i t m ill bo oboorrod th st tbs
sums gonorsj. trends fo r ssoh phsso sppssrod during tho f i r s t tmo hours
s f ts r i n i t i s l szposuro.

In ths eontlnuously trostsd group, mith ths

saosptiom of eam psrstirslj minor fluotustions, thors mors no rsdiosl
incrsssos or doorossss s f ts r tho i n i t i s l rosotion to Aoti-diono.

On

th is b ssis, tho tmo methods of trostmoat sopsmto snd i t msy bo ssid
th st the offoots during tho sntiro period of obsorrstion in tho oontinuouslj trostsd group sro oompsrsblo to those mhloh sppssrod during tho
f i r s t tmo hours of tho short trsstm snt.

Bsro, sgsin, ths most o ritio s l

ohsngss ooourrsd during th s ssrly hours s f t s r ths in itis tic n of li •stmsnti
In sunBsry, mith respect to tho hourly ohsngss in tho froquomsy of
tho individusl stsgos, tho follouing obsorrstions mors mods:
1.

The re su lts obssrvod in tho 1 ppi sorios mors mors simllor in

their gsasrsl trends to those found im tho throe lomost somoontrstiono

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of th# ahort tr##t*#Bt than thay «#r# to th# #ff##ta obaarvad im othar
eontianooaly traatad a a ta ria l#
S#

Xha raaa lta foumd im tha D ppa aariaa alaalatad thoaa ob­

aarvad im tha hi&har aoaaamtratiooa of tha abort traataaat im tha aarly
houra and, im tha la ta r houra, raaamblad othar aariaa im tha ooatimaooaly
axpoaad sroc^#
5#

Tha oatataadimg foataraa of tha highar aaaoamtratioaa (SO ppa,

40 ppa and 80 ppa) of tha eoatimnoaa aariaa aara tha deaimanoa of aataphaaa, tha aora or laaa a ta b ility of prophaaaa, tha groat daaroaaa im
poat-aataphaaa and tha ooaapiauoma laak of any radiaal ehamga a fta r tha
f l r a t hoar of axpoaura#
4#

Wham thoaa thraa highaat oomoantrationa ara aoaparad with tha

aama aariaa im tha ahort traatmant group, tha aama traada ara foumd to
ooour within tha f i r a t hour, but radioal ohangaa ara found in tha raaulta
of tha two typaa of tra a tarant in tha la ta r houra following i n i t i a l ax­
poaura.
5.

Tha moat o r itie a l ohangaa im a l l oomoantrationa ooourrad

ing tha f i r a t fiva houra a f ta r tha in itia tio n of traataaat#
Tha hourly variation# in tha fraquanelaa of tha individual atagaa
raaulting from tha two typaa of Intaxmittant traatmanta ara ahoam in
Taxt Flga. 86 and 87#

I t w ill ba obaarvad th at whan tha root tip# wara

axpoaad to a 40 ppm aolutiom for a pariod of 15 minuta# avary two houra
(Taxt Fig. 86) tha raaulta produoad wara alm ilar to thoaa obaarvad follow­
ing traatmant with tha aama aanaontration for a alngla axpoaura of tha
aama duration (Taxt Fig# 18)#

Tha only diffaramaa batwaan tha affaota

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of the two method# of epplieetion ### ea e a rlie r diaeppeerexiee of the
dlTleion figure# la the m ateriel that had beea la eoateet with Aotl—
dioae only oaoe#

In the merletem# expoeed to a 40 ppm eolation of the

aatlb io tlo for fifte e n mlaate# every hour (Text Mg# S7), the hourly
variation la the frequenolee of the Individual etegee mae# la It# general
aepeet, eimilar to thoee ohaagee obeerved la the material expoeed ooatla*
uouely to the eeme ooaeentratloa (Text Mg* SS)#
B# Specif le effeote of Aotl-dloae
The epeolflc effeote of the Aotl-dloae treatmeate were maalfeeted
in the aberrant figure#.

A deeorlptlon of them end a ooaelderatlon of

th e ir genetloal and cytologleal p o te a tla lltle e have prevloualy beea pre­
eented,

I t le of elgnlfloaaee to note that theee aberration# were not

found exoluelvely In treated material but oooaelonally did ooour (leee
than 10)1) In untreated root tip# (Appendix II# Table# 490# dSC# 7T0 and
900).
The hourly frequency peroentegee of theee aberration# throughout
the period of obeervatloa any be found la Appendix II# Table# 39 th ro u ^
90.

^om theee table# the data of the f l r e t five hour# were combined

(Table 17) and the peroeatagee of aberrant figure# la each phaae of
mltoele were calculated.
Text Mge# 70 through 7S.

Theee percentage# are graphically ehon In
By comparing the hletogreme I t la poeelble

to make enme general comparleone of the reeulte of treatment# for the
combined period of time on the three etagee of the mitotic cycle and#
In addition# to obtain eome Indication# of the elml l ar ltle e and d if f e r ­
ence# la the effect# o t the eeveral treatment# teeted#

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80
TABLE IT
mtOXNTAOBS OP ABERRANT PIODRBS OBTAINED
TROIE COMBINED DATA OP PIRST PIPE BOOHS

Prophaa*
COHO»*

T p # m tm # n t

Ahar. Total

%

Mataphaaa
Abar. Total
%

Anaphaaa
[. Total %

1 FRI

Short
Contlnuoua

63
515

1265
1468

5.0
35.1

9
112

547
584

1.6
20.4

7
12

231
188

3.0
6.4

5 F»

Short
Contlnuoua

635
1127

1479
1556

42.9
72.3

101
207

456
588

22.1
35.2

25
17

208
243

12.0
7.0

20 rMI

short
Contlnuoua

709
1022

1413
1378

49.6
74.2

188
374

430
923

43.7
40.5

36
22

256
106

14.1
20.8

Short
Contlnuoua
Short
Contlnuoua

622
620
995
587

1021
1133
1361
1187

60.9
54.7
73.1
49.5

lOl
493
132
287

424
911
480
888

23.8
47.5
27.5
30.7

43
44
32
9

177
121
233
62

24.3
36.4
13.7
14.5

70

1128

6.2

21

582

3.6

30

331

9.1

40 PIV
80 F »

untraatad
* Coneaatratloa

By rafaroaoa to tha thraa hlatogroaa (Taxt Plga# 70 through 7SJ a
eoaparlaoB of tha affaota of a l l traaW aata oa tha ihdlTldaal atagaa aay
ba aada.

I t w ill ba obaarvad th at tha paroaataga of abarraat prophaaa»

mataphaaa and anaphaaa flguraa waa h 1 |^ r In tha untraatad aarlaa t han i t
waa In tha Iowaa t aonaantration of tha ahort traatnant#

I t w ill ba aotad

alao th a t tha graataat fraqnanalaa of abarraat flguraa ooourrad in tha iro->
r**Taa group (Taxt Plga# 70 through 7S)#

Aa ahown In Tabla IT, with tha

azoaption of tha 1 ppm aarlaa, aora than 40% of tha prophaaaa wara affaotad
In a l l oonoaatratlona of aaoh typa of traatmant and. In aoma oaaaa tha
number axoaadad 70%#
^

Tha Iowaat paroantagaa of abarratloaa appaarad In

(Tbxt Pig. 72).

group wara balow 36.45L.

Aa ahom In Tabla IT a l l paroantagaa In thla
In addition, i t may ba obaarvad that tha graataat

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51
aiff#r#ne# between the two treatmente ooaeletently eppemre in the prophene hletogrem (Dnzt Mg. 70), while the meet elm ilerlty between the
peroentegee of eberrent figure# eppeer in the enephaee group (Text
Mg. 72).

M nelly, the 1 ppe ehort treatment mppeera to here been in­

effectual in inoreeeing the peroentegee of aberration in each of the
three etagee of mitoeie (Text Mge. 70 through 72).
▲oomparleon of the effeote of the eerered. oonoentratione and of
the two type# of treatment on each of the phaeee of mitoeie during the
oombined fiv e hour period
greme eeparately#

mey be made by a etudy of eaeh of the hieto-

Following the ehort treatment there wee a eteady in -

oreeee (42.9)1 - 75.1)&) im the frequeneiee of aberrant porophaeee (Text
Mg. 70) ae higher oonoentratione were weed.

Tbie differed from the

reeulte obtained in the oontinuouely treated group where, ae ehoun in
Table IV, the aberrant prophaae frequenoy dropped in the 40 ppm end
80 mm aeriee (19.9)( and

reepectively).

The greateet differenoee

between the effeote of the two type# of treahaent wore found in the
1 ppm and 8 ppm aete where the differenoee between the ahort and eontinueue applioationa ware 50*1$ and 29*4$ reepectively*

The 5 p]^ and

20 ppm oomtimuouely treated and the 80 pp^ 10 minute expoenre were the
moat effective ia producing aberration# in prophaee*
Following the ehort treatment there wae a rie e in the frequenoiee
of aberrant metaphaeee (Taxt Mg. 71) ae the concentrât ion wae inoreaeed
to 80 ppm when the percentage f e l l beak to the level i t had been at
5 ppm.

In the oontinuouely treated root tip e , however, the percentage

continued to inoreaee in the 40 ppa eolwtion, but dropped in the 80 p ^
tree Went*

The meet effective condition# fo r the production of

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■etaphesee were pverlAad by th# #0 p]pm «nd 40 ppm ##mtlmmo##
and th# 80 ppm ahort tr##tm#ata.

Th# p#ro#ot#@# dlff#r#n##a b#t###m th#

typaa of traatmanta la thaa# thra# aata of raaalta la probably not algnlfleant*

Tha gr aataat diffaramaa la affaatIvanaaa beta#an tha aama eoa-

eantratloaa of tha tao mathoda of appllaatloa appaarad la tha 40 pp&
aarlaa#
Tha moat a ffa a tlra traata&anta raaaltlag la abarraat anaphaaa# (Taxt
yig# 78) aara th a 40 ppa ooneaatratloaa of both typaa of traatmant#

Both

tha 1 ppi and 5 ppa aolutlona aay ba aooaldarad to taanra baan laaffaatlva
alaoa tha abarratloaa la tha traatad root tlp a dlffarad by #
from th a lr fraqmamay la tha oontrola#

or laaa

n ila dlffaranoa la probably mlthla

tha rang# of natural variation.
la aummary I t might ba aald th at from tha hlatograma aonatruatad
from tha eomblaad data of tha f l r a t flv a houra tha foUowiag pointa baaoma avidant:
1#

Tha graataat paroantagaa of abarraat flguraa raaulting from a l l

tha Aatl-dloaa traatmanta appaarad la tha prophaaa ataga.
8.

Tha graataat dlffaraaoaa batmaan tha aoma aonaantration of t

tmo typaa of traatmanta alao ooourrad la tha prophaaa ataga#
3#

Tha lomaat paroantagaa of abarratloaa ooourrad a t tha anaphaaa

4.

following tha 80 pjai ahort traatmant# tha raaulta produoad, la

ataga.

tama of fraquaaey of aberrant prophaaaa, mara almilar to thoaa appear­
ing la the 80 ppe and 40 ppe aoatlnuoualy traatad material.
5.

Tha 80 ppe aolutloa of both traatmanta aa mall aa tha 40 pjm

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■olatloo eœtinttously applied wmrm aqaally affaetlv# ia produe lac th#
hlgheet peroeatagee of aberraat metaphaeee#
6#

The 40 pj^ ooatlauoue treetmeat appear# to hare beea meet

effeotlTe in produolag dleorgeaiaed aaaphaeee#
The peroentegee of aberraat prophaeee are plotted in th e ir hourly
eequenoe la Text Flge# 28 th ro n g 41#

For purpoeee of comparleoa the

to ta l prophaee peroeatagee are iaoluded ae dotted line# ia theee gracie#
I t w ill be obeerred th a t in the untreated root tip e (Text Fige# 28 and
35) the peroentage of aberraat prophaeee reeutiaed ooaeieteatly belov the
10% level and the 1 ppm eoaoentration of the abort treatment (Text Fig#
29) wee v irtu a lly iaeffeetiv e ia iaoreaeing the iaoidenoe of th ie
aberration#
la the etudy of tho oomperative qu antitative effeote of Aotl-dloae
upon the frequenoiee of the individual etagee# one of the differenoee
obeerved betmeen the tuo type# of applioation uae the radioal hourly
variation in the prophaee end metaphaee frequenoiee following the ehort
treatment# while in the oontinuouely expoeed eeriee no eharp hourly
ohaagee ooourred a f te r the th ird hour and praotioally none a fte r the
f i f t h hour#

I f one eoane the graphe ehowing the hourly peroeatagee of

aberratioae in prophaee (Text Flge# 30 th r o u ^ 33# 37 through 39) end
alec in metaphaee (Text Fige# 44 through 47# 51 through 53)# one may
obeerre th a t the i r regular i t iee inoreaeed radloally during the f lr e t
f ir e hour# ae a re e u lt of both treatmente#

l a the la te r hour# of the

two lower oonoentratione In the oontinuouely expoeed group (Text Fige#
36# 37, 50# 51) divieion figure# had dieappeared and# ae a reeult# the

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54
graph# #r# mlallmr to thorn# r#pr###atlng th# ahort tromWomt#

During

th# ##m# porlod in hlghor oone#ntr#tlon# (Toxt Fig#, 37 through 39,
51 through 53) th#r# «##» ho##r#r, # gmdunl ri## In th# fr#qu#n#y of
nborrntlOB# In both prophm## and matapha##.
On# of th# ganaral affoot# o f Aotl«4Llon# «a# a daoraa## In th#
ralatlT# p#re#ntaga# of anaph### fIguraa#

On th# baal# of th# acmblnad

data of th# f i r a t flva houra (Taxt Fig, 7S), I t haa baan ahoun that
anaphaaa aaaaad to hara baan tha la a a t affaotad ataga#

From tha graph#

of tha hourly d la trlb u tlo n of paroantagaa of abarrant anaphaaa# (Taxt
Flga. 57 through 69) i t w ill ba obaarvad th a t tha paaVa of tha anaphaaa
dlaturbanoaa appaarad In tha la ta r houra of tha obaarvation#

In th la

ataga (Appandlx II# TOblaa 63 through 77C) tha para antag## ara banad am
aueh tmm 11#r nuabara and hana# tha vary radical ohangaa ara parbapa not
aa algnlfleant aa thay ara In tha graph# of tha othar phnaaa of mltoala.
Turning now to a aaparata oonaldaratlon of tha prophaaa abarratloaa,
I t a l l l ba raoallad th at ona of th a oharaotarlatlo affaota of tha ahort
traatnant nan tha ganaral Inoraaaa In tha fraquanay of tha prophaaaa#
Th# oouraa of th la inoraaaa la Indloatad by tha dottad lln a oa tha graphai
Omitting tha Inaffaotual 1 pjm aarlaa, whan ona aeana tha graph# of tha
raaulta follouing th# othar ahort traatmanta (Taxt Flga# 50 through S3)
ona obaarvaa a ganaral poa lt Iva and dlraot oorralatloo batmaan tha in­
oraaaa In tha paroantagaa of abarratloaa and tha Inoraaaa la tha fra quanalaa of tha to t a l prophaaa flguraa#

Tha 0 p]p&, 80 pjm and 40 p j^

aolutlona may ba oonaldarad to hava baan aqually affaotlva In tha aaxlnal
point# raaahad. In tha aohlavamant of thaa# navi ma in tha aarly houra

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06
a fte r exposure (three to four hours) end, follouing exposure to the tuo
louer oonoen tretien s of th is se rie s. In e dee lin e In the pereent of
eberretlons a f te r the peak haa heen r ess had.

The graph of the re su lts

obtained follouing a IS minute period In the 80 ppm solmtiom (Taxt Fig#
OS) stands alone In the series of thla method of applieatlom as haring
been moat effeetual slnee there uas 100% aberration a t six hours uhen
the to ta l prophase figures oen^lsed 75#9% of the to t a l division figures.
In other uords, slnee, a t the and of the period of obserratlon, the
dominant element in the observations uere these aberrant prophases, the
80 ppm se rie s differed from the re su lts found In root tip s exposed for
15 minutes to the other eomeentratioas#
When one observes the graphs of the data obtained from the eontln—
uonsly treated root tip s (Text Flga# 36 th ro n g 40), one Is Inellned to
oonelude that th is type of t r eatment uas more effective In produalng a
high pereent age of aberrations a t an e a rlie r hour than the Short tre a t­
ment method#

A olaser Inspection u i l l reveal one exception to thla: In

the 80 ppm aeries (Taxt Fig# 40) I t u lU be noted that the pereentaga of
aberrant prophaaes reached neither the peak achieved In the other con­
centrations of th is group nor the maxlmasi aehleved In the some eoaeentretion of the ahort treatment series#

fhrtharmere. I t may be noted that

th is general trend touar d an increaae In the percentage of aberrations
occurred regardless of the to ta l prophase pattern#
In the 1 PP& eons entretien of the continuously treated group
(Text Fig# 36) there appeared a tendeney for the to ta l prophaee percent­
age to Increase#

Thla ia ahoan by the dotted lin e in the graph#

It

u i l l also be observed th a t there uas a positive and direct correlatlcm

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56
between the Inereeee in nberrmtlon nnd the Inereeee In to ta l prophaeee*
In th le reepeet the 1 pi* eonoen trât Ion of thie group eorreepon&e to
the general tendenolee found in the ehort treetaent eeriee*

Beeeoee of

the rapidity of the Inereeee in the percentage of prophaee aberratioae
(95*4% in four houre)* the re eu lte of the 5 p%
m oomeentratlon of the
oontinuouely expoeed group (Text Mg* 37)* appear eu>et eimilar to thoee
found follouing the 80 ppm ehort treatment (94*9# in four howe)*
Ae the to ta l prophaee pereentagee deellned* the frequency of the
eberrent f Iguree Inoreaeed very markedly in root tip e eomtlnuouely ex­
poeed to the 80 ppm and 40 ppm eolutione (Text Tige* 38 and 39) and
henoe a d ire c t negative correlation uaa eetabllehed*

Ae a reeult* a t

the end of the twelve hour period of obeervation* the percentage of
aberrant figuree wae eloee to 100# but* a t thie time the prophaee etage
oompoeed between 40# and SO# of the to ta l divieion figuree and th ie may
be eaid to be an out et ending difference between the reeulte obeerved in
the two typee of treatment*
The 80 ppm eomtinuoue eeriee etande alone w ithin th ie group i a
having ehown radical hourly changée in the pereentagee of affected pro­
phaeee ae well ae im the fa c t th at the f r equency of the aberratioae doee
not reach the peak achieved in the lower concentrâtione of the eeme type
of treatment.
I t w ill be recalled that* on the baele of the frequency pereentagee
of the individual etagee* the reeulte of the interm ittent treatment a
appeared eimilar to the ehort end eomtlnuouely treated meteriale depend­
ing upon the length of the in terv al between the applicatione*

When the

hourly percentage of aberrant prophaeee ie examined (Text Flge* 34

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97
41)» ho##ver, both of th##* ##rl## la th# ##rly hour# #pp##r to h#v#
h#d #ff#et# mo#t hourly ulmllur to th# 60 ppm ooooohtrutlom of th# uhort
tr#utm#nt und, la th# lu to r hour# to bur# dlT#r##d la th o lr #ff#et#«
Whoa th# la to rru l hot ## on th# oocpoaur## uu# oa# hour uad 45 mlauto# th#
rooult# r#»#mbl#d tho## found la th# ahort trouteont aorloa, 1##., d l r l aloa figura# dlauppourod; uhoa th# la to rru l butuoua th# oxpoauro# uu#
only 49 mlaut##, th# raault# la th# la ta r hour# of th# obaorvutloa
porlod u#r# mor# hourly almilar to thoaa oboorrod la ocmtlauoualy axpoaad
root tip#» 1*0 . , th# d lrla lo a figura# poralatod u lth l l t t l # ohung# u a tll
th# and of th# alaraath hour.
Whoa cm# oxomlaa# th# graph# ahoulag th# hourly dlatrlbutlM i of
paroaatuga# of **#outt#r#d** matuphuaa# (Taxt Flga. 4£ through 59)» oa#
ahould f l r a t boom## uuur# of th# fraquaaol## of to ta l matuphuaa.

If

thaa# graph# ur# oomparad u lth thoaa of prophaaa (Taxt Flga. 88 through
41}» oaa u i l l obaarra th a t la thoaa oonoaatratloa# of tha ahort trauhaaat
aarlaa (Taxt Flga. 30 through 53) la uhloh abarraat prophaaa paroantagaa
uara r a la tl r a ly high thara uar# oomparatlraly high fraquanola# of to ta l
prophaaa flguraa a t tho aama tlma.

Ia tha aomparabla mata phaaa graph#

(Taxt Flga. 44 through 47) tha to ta l mataphaaa figura# daollaad ooaaidar*
ably #o that» by tha and of tha obaarratloa porlod» although 10O$ abarratlo a may bar# baan aaaa» th la partieu lar typa of affaot oaa not ba oomaldarad aa a dominant faatura of tha raault# of tha ahort traatmanta ulth
thaa# aolutlona.

Ia tha thraa hlghar oomoaatratloa# of tha eomtlauona

traataiant» tha to ta l mataphaaa fraquaaoy roaa» and» a t tha aama tlma»
tha paroaataga# of tha abarratloaa alao laoraaoad.

A# a raault» by tha

and of tha obaarratloa porlod» tha abarraat mataphaaa# may ba aald to

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se
hare b##n th# algnlfleant #lam#nta In th# r#aulta of th# oontlnuoiia
tr#atai#nt with th# hlgh#r oonoaatratlona.
In th# lowar oonoaatratlona of thla aarlaa, th# raaulta produoad
war# nor# naarlr almilar. In ganaral, to thoaa obtainad In tha ahort
traatnant aarlaa.

Although tha Inoraaaa In mataphaaa abarratloaa during

tha f l r a t flva houra waa mora gradual In tha group oontlnuoua ly axpoaad
to tha 1 pim aolutiom of Aotl-41ona Craxt Fig. 00), tha ganaral affaota
may ba aald to mora naarly raaambla thoaa axpoaad to tha 5 piai oonoantratlon for 15 mlnutaa during tha aama pariod (tUxt Fig. 44).

In tha la ta r

houra, whan th la fTaquanoy ahar ply daollnad In tha ahort traatmant
aarlaa. I t oontlnuad to rlaa In thoaa whloh ramalnad oomtlnuoualy In tha
Aotl-dlona aolutiom.

In thla raapaot, tha lowaat aonaantration of tha

oomtlnuoualy traatad group may ba aald to hava axhlbltad tha tandanolaa
of both typaa of traataiant oomaaoutlvaly.

On tha othar hand, on th la

baa la of Inoraaalng mataphaaa abarratloaa, tha raaulta found In tha
5 ppa aarlaa (Taxt Fig. 51) aloaaly raaambla thoaa notad In tha 80 ppm
ahort traataiant (Taxt Fig* 47) and othar oomtlnuoualy traatad root#
(Taxt Flga. SS and 85).

Howavar, tha to ta l mataphaaa fraquanay did mot

Inoraaaa grant ly and hamaa tha ralatlv a fraquanay a t tha and of tha obaarratlon pariod mora naarly raaambla a tha altuatlom found a fta r tha
ahort traatmant.
Tha dlatrlbutlon of tha hourly paroantagaa of ’*aaattarad'* mataphamaa ooomrrlng during tha Intarmittant traatmant ualng 40 pp* for 15
mlnutaa avary two houra (Taxt Fig. 48) oloaaly followad that obaarvad
following a alngla axpoaura to a aolutiom of tha aama oomoantrat 1om
(Tazt Fig. 44).

Whan tha Intarval waa ahortanad (Taxt Fig. 55), tha

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99
r#«ult#, during th# f i r s t f i r s hours, slanlstsd thoss obtnlnsd s f t s r m
single sxposttrs to the seme eoneentmtloo (Text Mg. 46), but in the
la te r hours the division figures remained in evldenee, a feature ehara o te rls tle of the 90 ppa and 40 pjai oontlnuoualy treated group (Text
Mgs* 59 and 55).
The following off sot# on anaphase have already been pointed out:
(1) One of the general e ffe c ts of the Aotl-dioae treatments was a réduc­
tion In the re la tiv e ntsbers of anaphase; (8) On the basis of the ooablned data of the f i r s t fiv e hours (Text Mg. 78), anaphase was the stage
le a s t affected by the hcti-dloae treatments; (3) A relativ ely high per­
centage of disorganised anaphases developed in the la te r hours, particu­
la rly in the root tip s continuously treated with the higher concentrations
of the a n tib io tic

(Text M gs. 66 through 68).

When the g r a tis of the hourly d istrib u tio n of disorganised anaphases
are examined (Text Mgs. 56 through 69) for a general comparison between
the effects of Aati-dlcne under various conditions, one finds relativ ely
the same general tendencies as those previously mentioned for the metaphase groups.

The following s im ila ritie s w ill be observed; (1) The 1 ppm

short treatment (Text, Mg. 97) waa v irtu a lly ineffective in disorganis­
ing anaphase; (2) During the f i r s t fiv e hours, the re s u lts of the 1 ppm
continuous treatments (Text Mg. 64) were most nearly similar to those
found In the same period In the 9 ppm short treatment series (Text Mg.
58), but In the la te r hours they simulated the re su lts obtained in con­
tinuously treated material;

(3) During the f i r s t five hours of soatiauous

treatment with the SO ppm and 40 p;m solutions (Text Mgs. 66 and 67),
the effec ts were somewhat similar to those observed in the s#me period

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60
followiac exposure to the seme ooneentretloas fo r the 15 minute period
but, in In ter hours, dlfferenoes appeered and rather radical changes in
the percentages of disorganised anaphases occurred; (4) In the in te r­
mittent treatment, mhen the interval betuesu exposures was long (l%xt
Fig* 62) the anaphase disorganisation formed a pattern similar to th at
found following the same concentration a f te r a single treatawmt (Text
Fig* 60).
In these same graphs the following differences between
and metaphaae e ffec ts can also be noted; (1) During the f i r s t five hours
in the 5 ppm continuously treated group (Text Fig* 65) there was a steady
ris e in the amount of anaphaee disorganisation but the level r eached did
not equal th at found in untreated material*

In the case of the metaphase

figures, on the other hand, th is solution was highly effective when con­
tinuously applied (Text Fig* 51); (2) The 80 ppm solution for the 15
minute period (Text Fig* 61) seems to have produced less disorganisation
in anaphase than occurred when the next lower concentration was used for
the same time (Text Fig* 60), a situatio n contrary to that found in the
metaphaee group (Text Figs* 46 and 47); (5) During the f i r s t five hours
of the intezm ittent tree Went when the roots were treated fo r 15 minutes
every hour (Text Fig* 69) the re su lts in tezms of anaphaee disorganisa­
tion were more nearly sim ilar to those found a f te r exposure to the 80
ppm solution for a single 15 minute period (Text Fig* 61)*

In the la te r

hours the aberrant anaphase percentages steadily increased in a manner
similar to th at observed when root tip s were continuously treated with
the 80 ppm solution (Text Fig* 69)*

These re s u lts more closely resembled

the effects of th is intezmittent treatment on the prophase stage (Text

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#1
rigm. 47, 94 mnA 59} than on m#tapha#* (T#zt Tig#. 39, 40 and 41}.
la Smeary:
1.

Tba 1 ppa aolatioa ahaa applied fo r a period of 16 alaute# aa#

iaeffeetlTa la produolng aberratloaa la aay of the a lto tla atagea.
3.

Oa the baais of the aomblaed data of the f l r a t five hour# the

high# at pereeatagea of aberratloaa appeared la prophaae aad the Loaeat
la anaphaee.
3.

The f l r a t five houra a f te r the In itia tio n of treatneat aaa the

period la ahleh the greateat pereeatagea of aberratloaa ear# prodneed la
prophaae and metaphaae.
4.

After the f l r a t five houra the moat radleal hourly variation la

aberrant aaaphaaea oeeurred.
9.

The greateat differeaeea betveen tha two typaa of treatnaat

appeared a f te r the f l r a t fiv e hours.
6.

During the f l r a t five houra of the la te z n ltte a t treateenta the

aberration pereeatagea reaembled thoae obtalaad f r om merlatena to ehleh
the amma eooeeatratloa eaa applied

a atngle time.

la the la te r houra

of the obaervatlon, the reanlta of these types of appUeatiom diverged
and, la thoae aerlea idiere the Interval beteeen expoaarea eaa short,
they ears similar to thoae obtained in the eontlauoualy treated material,
when the Interval eaa longer the re su lts simulated thoae found a fte r the
short treatments.
7.

Although the same general tendenolea eere ahem la the effeeta

of *11 ooaoentratloaa of the oontlauoua treatswnt, there appeared s drop
in pereentagea of aberrations In prophaae and metaphaae in the 80 p ^
aeries.

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68
In Appendix I I , 1%blem 78 throegh 90C the mmbere of prophase end
metepheee cell# eontelning "redootlonel" or "seeregetlonel" grouping#
ere presented in th e ir hourly eequenoe together with the peroentege they
occupy of the to te l of the propheee end metepheee dlvlelone.

rrom the

peroentege# obtelned during the f i r s t five hours the meen end I ts atenderd error mere celculeted.
shoen in Text 7ig« 73#

These everege percenteges ere grephlcelly

In th is figure I t #111 be seen thet the 40 p%
m

solution of both methods of treetment eppeered to here been most e ffec t1rs In Increesing the frequency of these types of eberretions#

Is

Teble V these everege percenteges ere presented together with th eir
stendsrd errors#

Here I t mey be observed thet the meen peroentege ob­

tained following the 40 jrpsi short end in the 80 pgm, 40 ppm end 80 ppm
continuous treatments were found to have been s te tls tI c e lly significant#
TABI8 T
M
EAN PERCENTAGES OY FROFHASX AM) METAPHASE CELLS OONBLININO
«REDOOTIGNAL** AND "SEGREGATIONAI," OROUPINQS

Conee n trâtion
1 pm

s pm
80 pm
40

pm

80

pm

Untreated

TYeetment
Short
Ocmtlnuous
Short
Continuous
Short
Continuous
Short
Continuous
Short
Continuous

Meen %

Stemierd Error

1.1
1#8
8.17
3.11
£#45
354
5.58
6.10
£.46
5.48

t 0.15
- 0#51
X 0.56
- 0.64
- 0#60
t 0.89»
2 0.68»
t 0.63»
X 0.66
t 1.48

1.80

t, 0.44

* S te tls tic e lly significant a t the 9^ level

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DISCUSSION
A« G#m#r*l

of Aoti-^iono

On th is bonis o il of tho doto ooovomlotod in this oorioo of
ozporlmont# ono aoy vemtur# tho hypothool# thot tho gomorol offoot
of tho Aotl-dlGoo trootmonto oppoorod to hovo boon #1mllor to thot
of o oloo fixing ogont.

Whon the doto oro o r i l nod in the light of

this hypotheoie the reoulto observed boeomo underetondoble.
Whon the oreroge mmboro of divleion figoree were oxeminod,
eontrootlng trends «ore found to ohorootoriso the too moin types of
▲eti^ione trootmonts studied in th is Investigotion*

follooing tho

short trootmonts thoro oos o gonorol doe lino in tho overage snmbors
of dividing s o ils , obilo in tho so root tip s eontinuonoly exposed tho
ulttmmto offset oos o s ta b ility in frequoney of division figures*
In the short treotments* in «hieh tho root tip s «ere exposed to Aoti*
diono fo r 15 minutes, the ossumod fixation effect «os ineomploto end
«hen removed from the to st solution tho s e lls «ere s t i l l eopoblo of
reverting to o "resting** condition*

Oonsequontly, there «ere no divi­

sion figures in the la te r hours of tho obeorvstion period*

In those

root tip s continuously exposed to the more effective eoncentrotions of
Acti-^ione, however, tho nimbors of figures a t tho end of tho obeervetion period «ere approxlmotoly tho some os they hod boon at tho time
treatment «as initiated*

Furthermore, a fte r tho f i r s t five hours

(Text Figs* 11 and 1£) there «oe l i t t l e change in the average niscbors
of dividing cells*

On the basis of th is evidence i t appears thot, in

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64
th### ooDoentr#tlon#, th# o#U# v irtu a lly b#oom# flz«d aftar a parlod
of five or six hours.

TIm amount of Aoti-dions in ths lower oonosntra-

tlona of tha eontinuoua treatment (Text Figs. 9 and 10} also appeared
to have been ineuffioient to have eoapleted the aeew&ed fixation and,
as a re su lt, by the end of eight hours tha e e lls had reverted to a
"resting" condition,

un tha other end of the concentration series, i . e . .

In the highest concentration (Text Fig. 13), the amount of Acti-dioma
probably had r sac had a levai mhera tha fixing effect mas nearly completed
to a point ju st below le th a lity .
)#hen tha re lativ e frequencies of individual stages mas examined
th is hypothesis mas corroborated (Text Figs. 14 through 87).

Following

exposure to the short treatments, there mas a more or less general rise
in the re lativ e frequency of the prophases.

In the early hours th is

ris e mas inversely proportional to tha concentration, while in the la te r
hours the relationship mas almost a d irect one.

I f me assume the same

numbers of c e lls may enter and leave each phase then, with Acti-dione
operating as a slow fixing agent this relationship i s to be expected.
In the early hours following the lower concentrations of both treatments
(Text Figs. 15, 16, IT and 80) tha fixing action mas quite weak and very
slow resulting In an accumulation of prophase, while in higher concentra­
tions (Text Figs. 18, 19 , 88 , 83, 84, 86 and 87) i t appeared that the
sta llin g effect on the changes had been increased to the extent that the
numbers of c e lls passing into prophase mere being affected, thus, with
fewer c e lls coming into division, and those present being slowly fixed
an apparent e ta b ility mas reflected.

I t appeared that the cells mere

not completely fixed in lower concentrations with Acti-dione but, as

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65
praTioualy Ixidloatad» rarartad to iBtarphaaaa*

Latar ataga# of tbaaa

rararaiona hava tha gaaaral aapaota of "affaotad" prophaaa naelai*
aooordlogly, tha rla a in prophaaaa a t tha and of tha pariod of obaarvation (Taxt flg a . 15 through 50) appaarad to hava baan a raflaetlon of
thla rararalon and an aacplanation of tha d iraat oorralation batvaan aoneantratlon and fraquanoj of prophaaa following abort traatmant#

On tha

othar hand, la thoaa root tip# eontlauoualy axpoaad to higher eoneantratlona, tha fixing affaet arantually aaamad to have baan nearly eonplatad
(probably at tha and of about fiv e houra) and tha raau lt waa a relativ e
a ta b lllty In tha fraquaney of th la ataga (Taxt Plga. 22, 23, 24 and 27)
throughout tha raaaalnlng pariod of tha obaervatlon*
I f wa aaaima tha alow a ta llln g a ffa e t In prophaaa aa eaualng tha
Inoraaaa In tha relativ e numbara of thla ataga, a daeraaae In tha relative
fraquanclaa of la ta r atagaa of mltoala would be axpaetad*

Tha data ahow

thla to have baan tha eaaa (Taxt Riga. 15, 16, 17 and 20) In tha early
houra of and following traatmante with tha lower eoneantratlona of Aetldlona.

On tha othar hand. I f , In tha higher eoneantratlona, there la

auffie lent Aetl-dlona praaant to have not only langthanad tha duration
of tha atagaa, but alao to have Inoraaaed In a ffaet to tha extant that
tha raau lt la a lowering of tha nuabar of c e lle cowing Into dlvlalon,
and I f wa alao aaawa that tha a ta llln g a ffect on tha aavaral atagaa la
a d iffe re n tia l one, than whichever ataga la laaat affected would be axpaoted to ahow a decline In relativ e frequency*

Praaoaably, therefore,

an increaae In tha re la tiv e frequency of a given atage at a given tla a
would Indicate that I t la the one moat highly affected by tba atalllng
action a t tlw mowant.

For example. In tha f l r a t three houra following

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66
the ehort tremtmeot with the 40 ppw eolation, metepheee eppeered to here
been the most effected*

During the eerly hours of the obserretion

period th is wes found to here been the oese In the root tips exposed to
e l l the higher oonoentretlone (Text Tigs. 18, 22, 23, 26 end 27) idiere
there wes e sufficient amount of Acti-dione present to here increesed
the Intensity of the ste llin g eetion to th is extent*

Ths 15 minute

tree Usent wes epperently not long enough for e complete fixation end,
es e re s u lt, depending upon the degree of the e ffe c t, dividing cells
could e ith e r complete their division or revert to interpbese*

In the

continuous treetment, however, the c e lls , being almost completely fixed
by the end of four hours, there Is , es would be expected, l i t t l e chsnge
in the re la tiv e frequency of the individuel stages a fte r this time*
I f the graphs of e ll treatments (Text Tigs* 13 through 87) ere
divided et the point representing the end of the f i f t h hour I t w ill be
observed th et the relative frequency of post-mete^ieses declined In the
f i r s t hours considerably end, depending upon whether or not the influence
of the Acti-dione bed been greet enough to completely **flx” the c e lls,
the two types of treets&ent diverge e t th is point*

In short tree Went

series (Text Figs* 19 through 19) end the lowest concentration of the
continuously exposed group, epperently only e slig h t fixing of the cells
bed oocured*

In the other series of continuous treetment, however, the

poetHsetephese c e lls appear to hove been almost completely fixed with the
re su lt th e t there wes l i t t l e change in th e ir re lativ e frequency after the
f i r s t five hours*
The resu lts of the interm ittent treatments also support this

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67
hypothe#!# of » fix a tio n action of Aoti-diona*

In came# ahara tha

root tip a aara axpoaad to a 40 ppa aolution of Aoti-dioca for 15
ainutaa arary tao hour#, during tha f l r a t fir a hour# tha ralatlv a f r a quaneia# of tha individual ataga# aara aimilar to thoaa obaarvad ahan
tha aama cone e n trât ion aa# uaad for a aingla 15 ainuta pariod.

Aftar

fiva houra, tha diviaiona diaappaarad in tha l a t t a r aariaa, but in tha
in tax n ittan tly treated m aterial tha rarvaraiona appear to have taken a
little

longer and diviaion figure# remained in aridanoa fo r an additional

thraa houra#

Whan tha in te rv al bat uaan tha traatmanta mas ahortanad to

45 ainutaa tha raault# obtained (Taxt Fig# 27) nara aim ilar to thoaa
obaarvad ahan tha root# vara oontinuoualy axpoaad to tha aaoaa aonaantration#
In atnmary, tha data axnmlaad ao f a r aupport tha naaoaption that
Aoti-diona haa a d iffe re n tia l alou fix atio n a ffec t ahich, in loner con­
centration#, raault# in lengthening of prophaaa and nataphaaa and In
high concentration# th ia a ffe c t incraaaaa raaulting alao in lowering tha
number# of call# coming into and going through thaaa diviaion figure#.
B#

Tha Specific Xffacta of Acti-dione

Tha typaa of abarrationa. th e ir intarralationahipa and alnnif icancaa#
Tha more apacific affect# of Actl-diona may be conaidarad to have baan
raflaotad in tha varioua aberration# th a t ware obaarvad#

In tha jraaaat-

atlon of tha raault# tha many typaa of aberrant figure# ware daacribad.
Basantia lly tha irra g u la ritia a reported are aimilar to acme of thoaa
praviounly obaarvad by othar invaatigatora (Appendix I)#

ïha practical

importance of tha c ample ta ly diaorganixad figure# (c-a&itoaia) i* wall

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68
knomi (18, 45, 47, SO, 70, 79, 99 and 144)•

Sana of tha aora thaoratieal

p o ta n tla litia a of tha organlmad **rad\iotioBal** grouping# bara baan pra—
Tlousljr diacussad (62, 89 and 142).

All th is work «as baaad upon eon-

tlnuouslj traatad n a ta ria l obaarvad a t varying intarvals ranging from
four to taalva houra*
In th is study tha affaet# of a 15 minuta troatamat pariod «era
eontraatad «1th thoaa oontinuoualy axpoaad and tba quantitative data
«ara eoUaetad a t hourly in ta rv a ls.

Baaad upon tha re s u lts of thia

study, a general aunary of tha typaa of affaota, possible in ta rra la tionahipa and tha p o ta n tla litia a of each i s praaantad la Table VI.

In

th ia table i t « i l l be seen th a t, depending upon «hathar or not there i s
an element of organisation praaant, the *raduetional" groupings (Plata
I I ) may be separated from tha "asgregetional" type (Plata I I I ) and tba
"seattarad" arrangemanta (Plata TV).
Although tha "redaetional** groupings are primarily of intaraat
baeauaa of tha p o ssib ility of th a ir resultin g in a raduetion in tha
ehromosoma number « ith tha eonaaquenea, in many eases, of providing for
genetieal segregation (62), othar speoulations on th a ir further develop­
ment are ahomn in Table VI.

As previously mentioned, either equal or un­

equal numbers of ehromosoma# may be involved in these "raduetional* group­
ings.

When, in hataroaygous stock, homologues are separated in thia

manner, thaaa abarrationa aaanaa ganetioal aignifloanee.

In thoae oases

of prophaae and metaiAase *reduetional* groupings «here equal nWbers
are involved (PI. I I , Fig. A, also Prontispieoe Pig. A), the daughter
ohromatida « ith in eaoh group must be separated into t«o individual
nuolei i f the original number is not to be restored.

This would resu lt

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CD

■
D
O
Q.
C

g
Q.

TABXJ VI

■
D

CTTOLOGICAL AMD GEDBTXCAL P O T lM T IA L m iS 0 7 ΠB SEVSHAL TYPES OF ABBffiATIOMS

CD

C/)
C/)

NORMAL
8D
■
CYT0L06ICAL

EFFECTS

rTOXICITY EFFECTS
1

CD

3.
3"
CD

CD

■
o
O
Q.

ORGANIZED FIGURES

rREDUCTIONAL
GROUPINGS

DISORGANIZED FIGURES
SEGREGATIONAL"
GROUPINGS

o

NUCLEAR MEMBRANE

KCHROMOSOME STICKINESS-i
-"SCATTERED"!
(GAP 0PENS)ARRANGEMENTS
— - T - ^
T
REVERSIONS^
(GAP
CLOSES)

C

a
O
3
"O

1

n
^EOUAL
NUMBERS-

i-NUM BERS

r ~
EQUAL

unequal

NUMBERS-

1

UNEQUAL

NUMBERS

-►DETERIORATION OF
NUCLEAR CONSTITUENTS

CD

Q.
P L IT
lT RETAINED
REl
rGAP C LO
OSSEEsSJ^^S
SP

SPLIT RETAINEOJ#GAP a O S E S

■
D
CD

(/)
C/)

RETURN

TO
DIVISION
PATTERN

-P GENE TIC SEGREGATION

^GENETIC SEGREGATION

normal

\

h R oÏ oV Ô m E
CHROMOSOME

no

NO

REDUCTION
CHROMOSOME NO

ANEUPLOIDY

POLYPLOIDY-4.

70
In th# produetloa o f four auoloi ##oh eoataialnc olght ohromomom###

If

thlm ohouU ooour, gonotle oogreg&tloa and roduotioa in ohrcmomcm# numb­
er# would be neeompllebed e# shown In Table 71.

One» perhaps question­

able» case of th is type of separation is shown in f t^ tis p ie o e Tig. D»
where four groups of eight chromatids have been formed in the seme c e ll.
Here the likelihood of the formation of a tetrad of four nuclei each with
eight chromosomes s a i s highly probable.

However» none of the few re­

duced c e lls found during the course of th is study contained eight chromo­
somes.

I t sctms logical» therefore» that» the actual re su lt following

these "reductional** groupings may be e ith e r an elimination of the sepa­
ratin g gap and a return to the normal condition, or the formation of
nuclei with the reduced chromosome mmtber which» fo r the most part» can­
not divide again.

Because of the element of organisation in the figure»

the return to the normal condition may be affected in the stage follow­
ing that in which the separation occurred.

Tor example» the chromosomes

composing the two groups in TTontispiece Tig. A may close the gap between
them during the course of th e ir movement to the metaphase plate and a
normally organised metaphaae figure might result#

I t seems reasonable

th a t the return to the norwml division pattern would depend upon tha
orientation.
As shown in Table 71 when the prophase and metaphase separations
are unequal the same two p o s s ib ilitie s present themselves:

Either they

may return to the normal division pattern in the succeeding stage or
the gap aay be retained u n til the division ceases*

I f the la tte r occurs

there are two further probabilities» both of which lead to aneuploidy#
Each group of whole chromosomes may form a separate nucleus.

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Tor

71
example, i f the two g r o u p » in the 10*6 eeperetion ahomn in P I, I I ,
P is, B were to form aeperete nuolei end the oonatituent ehrometida
aheuld f e l l epert in a itn the re a o lt would be two nuolei with 80
18 ohrcaneaomoa reapeotively.

On the other hend, i f the ohroamtid# of

the 10*6 "reduotionel** groupins ahould be aepereted into nuoloi, the
4 deughter nuolei would oontein 10, 10, 6 end 6 ohromosoawa reapeot*
Ively,
Some evidence aubetentieting the retention of the gap in ceaeo
involving aeperetiona of unequel ohromoaome numbera aey be found in
the oooeaional obaervetiona of unbalanced telopheaea, of binucleete
end trinucleete c e lle with unequal aimed nuolei end, in the very rare
appearance, of reduced c e lle .

But i t ia not poaaibla to daoido whothar

thoy aroae in th la manner, from more or leaa organ!med fignroa, or from
the completely diaorganimed type.

The point to be atreaaed here ia

th a t, although the evidence of actual reduction waa rare in thia atudy,
the genetieal aignifioance of the "reductional* group ia , nemethaleaa,
not diminiahed,
Aa ahomn in Table 71 two typaa of d iaorganimed fignroa wore obaerved.

Thoae have been deacribed in the vreaentation of the reanlta

and ere deaignated aa the "aegregetional* gronpinga and the "acattered*
arrangemanta.

In the previona lite r a tu re theae figure a collectively

are referred to aa **CHaitotic** diviaiona.

The "aegregational* gronpinga

have been called by varioua terme, "exploded metaphaae**

( 6 ),

trlbuted o-mitoaia* (106) and "diatribntive c-mitoaia" (85).

"dieAa ahown

in Table 71 theae "aegrcgational** gronpinga may involve the aeparation
of eith er equal or unequal nim#bera of chromoaca&ea.

When equal numbera

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7M
of ohromoocm## haro b##n mepmrated thay ara of partioular Intaraat ainaa
thay rapraaaat tha poaaiblllty of raaulting in ganatieal aagragation
wlthont Involving a raduotlon la ofaroaoaoaia ntmbar. Vhan two mmarloally
unaqual groupa ara aaparatad, juat aa In tha oaaa of tha "raduatlonal"
grouplnga, anauploldy may raault#

A ltam atlvaly, howwva r # thara la tha

additional p o aaib lllty of tha Intargroup gap oloalng and raaulting In
"aoattarad" arrangnaanta, whleh may avantually land to polyploidy.
Tha aaoond typa of dlaorganlsad flguraa (Saa Tabla VI, paga 69)
la th a t uhioh hara haa baan daalgnatad aa *aoattarad" arrangmmanta*
Thaaa wara iU u atratad In P lata IV and In Flrontlaplaoa Plga. S and P.
Thair potantlal oapaelty for raau ltin g In tha polyploid aondltlon la
wall raoognlsad but I t might ba of In taraat to point out, aa Indieatad
by dottad arrowa Tabla VI, th a t, onoa aatabUahad by a **drlftlng" of
whola ohrcmoaomaa, thaaa "aoattarad** arrangomanta may. In aona eaaaa
glva r la a to "aagragatlooal" gronpi nga.

In th la raapaat thay alao pro—

aant a poaaiblllty of raaulting in alth ar anauploldy or ganatio aagra­
gation without raduotlon In ohrcmoaona numbar.

I t waa polntad out, th at.

Ilka tha "aagragatlonal" groupa, thaaa "aeattarad** arrangmmanta wara
alao moat eharaotarlatia of tha l a t t a r atagaa of oontlauoua traatmant
or tha l a t t a r houra a fta r axpoaura to tha ahort traatmant and appaarad
to r a f la a t tha eomplata Inhibition of tha aplndla.
In aunmary of tha oytoganatleal p o ta n tla litia a of thaaa abarra­
tiona:
1.

Tha ultimata ra a u lt of tha oraanlaad diviaion flguraa may ba:

(a) a raturn to tha normal diviaion pattam ,

(b) ganatio aagragation

with raduotlon In ohromoaoma mnbara, and (o )anauploldy.

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▼
9
£•

The dleor«eni»ed eomflgeretleam aey lead to:

(e) gene tie

segregetiOA witheat réduction ia ehroaoeome aœbere» (b) eaei^loidyt and
(e)

polyploidy*
The eytologioal aberration#, th e ir re la tiv e freauenoiee end relm-

tionahipe to tm ee of treatment.

TP diaoorer any poesible relationehipe

beta#en the freqpanoie# of theae aberration# and the aonaantration, the
tine aa well aa the method of treatment uaa one of the more apeaifia ob—
jaativaa of th la in raatig atio n .

I t maa hoped th at i t mould ba poaaibla

to obtain a apeaifia oombinatioa of theae factor a uhiah mould raault in
the product ion of one or more reoorarable abarrationa and i f th ia mare
not poaaibla to define the threaholda of aytologioal and tonie effeata.
In the dieouaaion of the generel effeata of treatment i t maa
pointed out th a t the meat o r itic a l period fo r the obaerration of the
eytologioal effeeta of Aeti-dione maa the f i r a t five houra a fte r the
in itia tio n of treatment.

Thia early in terv al had not been atudied quan­

tita tiv e ly prior to thia inveatigation.

In the atudy of the hourly var­

iation in the frequency of diviaion figura# i t maa ahoun th at during
th ia time the greateat changea in re la tiv e nwmbera of diviaion figure#
oeeurred.

A alom atallln g action» d iffe re n tia l upon the individual

atagaa maa poatulated.

I t ia conaidered to be of aignifioance that»

in term# of abarrationa alao» there maa a d iffe re n tia l quantitative
e ffe c t.

Thia waa ahomn in Taxt Figa. 70 through 72 mhere the percent-

agaa of aberratione from the combined data of the f i r a t five houra were
graphically preaented.

Here i t maa indicated that the h igheat percent—

agaa of abarrationa occurred in prophaae and the lomeat in anaphaee.

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74
I t w ill b# ob##rved, th at, tha ralationahip of the degree of e ffect on
the re la tiv e frequeneiea of Indlrldaal etagea with the eoaeeatratlotta
and the type of treatment la not the aame aa that exlatlng between the
pereentagea of aberratloaa and theae varlablea.

On the baala th at the

relationahlp doea exlat one may poatnlate th a t the aaeumed a tallln g
action may be the preenreor of the production of the abarrationa.
In the quantitative atudy of the eytologioal abarrationa the
f l r a t five houra alao appeared to be the moat c r i t i c a l period.

Follow­

ing the abort treatment there waa a d irect correlation between fre ­
queneiea of diviaion flgurea and pereentagea of abarrationa in prophaae
(Text Figa. £9 through 33), but an Inveree relationahlp In metaphaae
(Text Figa. 44 through 47) and in anaphaee (Text Figa. 58 through 81)*
In a l l concentrationa of the oontinuoualy treated root tlp a the aberrant
diviaiona in a l l mitotic phaaea continued to riae even a fte r a a ta b lllty
in numbera of diviaion figurea had been reached, (Text Figa. 38 through
40, SO through 54, 84 through 88).
Furthermore, i t w ill be obaerved from the graphe th at the pereent­
agea of aberrationa were not in d ire c t proportion to the concentration,
but rather were more cloaely correlated with time and method of tre a t­
ment.

I t ia of in te re a t to note th a t the peak of prophaae and metaphaae

aberrationa in the 80 ppa of eontinuoua treatment waa lower than that
obaerved in more d ilu te concentrationa need In the aame manner.

Further­

more i t ia conaidered ecaKewhat aignificant th a t, in terma of pereentagea
of aberrationa the 1 ppa ahort treatment (Text Figa. £9 , 45 , 57) waa
v irtu a lly in effectiv e.

On the baala of the re la tiv e frequeneiea of the

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78
iadiridual

the 1 ppe short treetment eppeered to here been

effective In the In ter hoars.
I t seems reesonsble to eonolude from th is thet Aeti-dione, even
in th is d ilu tio n , hed oeused some disturbenoes in the regulsr eetivi t l e s e ith ln the nuoleus th e t resulted in the eeeimuletion of propheses*
Since t hese divisions could, be considered to heve been within the renge
of Bozmelitj, the effect wes not su fficien t to cease e breekdown of the
process but merely seems to heve hed the s te llin g effect mentioned ebove,
All of these re s u lts tend to support the hypothesis msde e t the
beginning of th is discussion thet the Acti-dione eppeers to not in e
manner simiier to e slow fixing egent.

I t eppeers th e t, es A ctivions

pénétré tes the dividing c e l l , i t s f i r s t e ffec ts ere slig h t physiologicel
disturbenoes which menifest themselves in two easily detectable ways:

,

There eppeered to heve been e slowing down of the ra te of some

1

of the morphological changes thet generally occur.

One of these activ­

i t i e s thet se#med to heve been effected wes related to the nos leer nsmbrene.

I t eppeered to heve been retained es e membrane end the resu lt

wes the eppeerences of the "effected" propheses (PI, V, Figs, B, C end
D),

In these cases the ehromoscmes themselves seemed to heve been

cspeble of proceeding through th e ir usuel s tru c tu re l development toward
metepheee while the nucleer membrane did not change,

%e resu lt wes on-

oriented chromosomes of metepheee length enclosed within the nucleer
membrane,
2,

There seemed to heve been e rath er slow end graduel in terfer­

ence with the spindle mechanism.

The p a rtie l inhibition manifested i t ­

se lf in the "redactioaael" groupings which appeared in the early hours

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76
of oboorrmtiOB maA tho ooaploto enpipreeslon woo lodlooted by the
"eegregotlonol" end "seottered** orreagmmemte ehioh eppeered in th eir
greeteet frequenelee la the le te r hoare.
Hone of the ooneemtretloae need la the abort treetment eerie#
of th le lareetlgetloa eeomed to be eafflelen tly high to here ecmpletely
fixed the e e lle during the 15 mlaate la te r r e l, henee, uhea remoeed from
the aolution the rerereele to renting etege eould oeeur*
from e l l treeteente Including the eontlauooaly treated group
there uea reeorery In only too of the eeriest the 1 ppm end 5 ppm ahort
treetment#.

The 1 ppm me# found to here been eytologleelly ineffeetlve

(Text Figs. 29, 45 end 57} •

Homsrer, la terms of the r e le tlr e fre­

queneiea of the ladlrlduel steges (Text Fig# 15), es prerlously pointed
out, acme effeeta mere mealfeated in the eeetaraletlom of propheses ead
the drop In r e l e t l r e frequeneiea of post-propheee steges mhieh oeeurred
a fte r f ir e hours*

%e eomperet 1rs ly normal d lrlslo n s la the reeorered

materiel Indleete thet the eheages Inrolred la the ste llin g eetion of
Aetl-d.lone mey be preperetery to the eytologleel eberretlon#

Further­

more, sines there me# no erldenee of say undue eytologleel eberretlon
In the reeorery materiel a fte r the 5 ppi short treetment. I t semes
reeeoaeble to esswme th e t the e e lls obserred In th is reeorery materiel
hed either eseeped the effeeta of Aetl-dlone or, at le a st, hed not been
redieelly ebanged by I t .

Slnee the 1 ppm short tree Went mas eytolog­

le e lly in e ffee tlre but did re fle e t some eheages boring oeeurred end the
5 ppm of the seme group prodused eytologleel o ff sets but appeared to
here been toxle to the c e lls eo effected. I t seems logloel to assume
thet the to x le lty threshold Is rery close to thet required for ths

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T?
produDtioB or eytologioal abem tiono*

I f rooorerablo aborratioae are

to bo obtainod from Aetl-dlomo I t oeome moot llkoly that thoy mould bo
found In oemo oombinatioa of timo from botmoon 0 and 15 miautoa and at
oomo oomeontration botmoon 1 ppm and 5 ppm.

At any rata i t

highly

probablo that by further inrootigation of th ia rango of troaWont, tho
amomoro might bo found to oomo vary portinont quostiono rolatod to tho
method of broekdoum, tho rooororability of aberrationa and poaaibly the
meohenima# involved.
The queotlon that next arioeo io , mhy i t io that tho oolla in
whioh Aoti-diono haa boon aetivo havo Loot th e ir a b ility to divide.
Tho only oboorvationa uhioh might poaaibly oontein aomo oluoa in anamor
to thia queation moro tho naturo of tho interphaaoa.

After expoauro to

tho a n tib io tio , tho interphaae nualol aaaumed a d iffe r ont appearanoo.
Thia maa partioular ly notiooablo in tho higher oonoentrât iona of both
typoa of treatment and, of oourao, maa moat aooontuatod in t hoae fixed
a f te r long porioda of eontinuoua troatment.
theae deteriorated intorphaaoa are ahowi.

In Plate VI, Fig. A tmo of
Aa provioualy montionod, i t

mao obaerved that thoro aoiod to havo boon a earn intorforonao with the
ooiling end the nuoleua had tho general appearanoo of a Zygotene.

Mom,

i t io moll rooognisod that in tho oaao of tho llluatratod ooU thia maa
moot likely a toxioity offeot.

Bomover, aimoe none of tho oolla mhioh

had menifoatod eytologioal of foe to of Aoti-diono more oapabla of later
mitoooa

oinoo di erupt ion of tho apindlo moobanimn in one diviaion

doe# not neoooaarily inhibit auooooding diviaiona, i t aewa roaaonable
that intorphaaoa are alao vulnerable to the Aoti-diono.
In thla roapoot tho motion of Aoti-diono différa from that of

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78
oolehlolM and giraa support to th# oontonticms of Oallumky (58) Allan,
Wilson and Powall (8) Loralass and Rarall (91) and otbsrs, uho oontanA
th at a ll ohomioals oapabla of produelng tha o-amltotlo affaot do not
naoassarily oparata In tha m
ama mannar.

I t i s opposad to tha opinion

of Laran and Lotfy (65) who appaar to ba working on tha altam ata aaaxmptio n .

Thay mantion (85) that in substanoas, "such as sodim nuolaata

which in strong oonoantrations glva an ineostplata o-SKltosis, a l l kinds
of daviating typas of oHsitosis and intaimadiata stagas batwaan 0- and
normal ara fraquant."

Accordingly, thay consider tha prophaaa "raduc-

tional** groupings as **daviating** typas and suggest that thay ara prophaaa
in nuclei resu ltin g from preceding "axplodad oHsitosis** or **distributiTa
c-mitosis.**
ITom tha re s u lts observed in th is study tha Lavan-totfy eonoapt
cannot ba applied In tha case of Actl-diona#

F ir s t of a l l , thaaa "sag-

ragational** groupings occurred most generally In tha la ta r pariod of
observations and in those series where thay occurred in gr eatest fre ­
quency none of tha c a lls racovarad#

In those cases where recovery did

occur ths "sagragational** groupings wara ra re .

In tha 5 ppm short tra a t­

mant ear la s , fo r instance, only 6 of tha 47 chromosomal groupings wara of
tha **sagragat ional** type#

Tha remaining 41 of tha **raduotlooal** nature

eould not possibly have arisen from preceding **azplodad c^titosis** since
thaaa disorganised figures do not occur In such frequency in untreated
material#
une of tha significant points revealed by th is study was tha
apparent correlation between tha time and tha frequency of tha aberra­
tion of a partioular type#

I t has baan pointed out that tha "raductionaJ

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79
«roupiasa of tha orsaniaad type ooetnxed la higheat faeqaeaelea In tha
early houra a f t e r In itia tio n of tree Went and th a t the more dlaorganlaed flgurea mere found In largeat numbera la t e r .

Thla mould aeum to

aupport tha hjpotheala th a t Aetl-dlone aeta in a —

to that

of a alom fir in g agent and glrea a further clue aa to the reaaon for
the meekneaa In the effect*

I t appeera lik e ly th a t Aetl-dlone pane

tra te a the c e ll very alomly and reanlta In a aequenoe of event a:
1.

A blocking or a ta llln g of the uaual nucleer a e tlv ltle a

before any aberration appear a#

Thla maa manlfeated In the 1 ppa ahort

treatment where there maa a change In the re la tiv e frequeneiea of the
Individual atagaa a fte r the f l r a t fiv e houra but no undue aberration.
S.

An apparent delay in the uaual a c tiv ity of the nuclear mem­

brane without an a ffe c t on the nexmal changea of the chromoaomea.

Thia

waa ahocB In the early appearancea of the "affected" prophaaea which
contained chromoaomea th a t appeared to have been capable of paaalng
through th e ir normal morphological changea p rior to metaphaae.

In

theae caaea I t would aeem th at Acti-dione e ith e r bad not r eached the
In te rio r of the nuoleua owing to the alow penetration or maa Ineffective
upon the chromoaomea thamaelvea*
S.

A delay in the formation of whatever atructurea bad not been

formed a f te r auffioient Actl-41one haa accimmlated to a ta rt "fixing" the
oonatltuenta of the cella#

Thla la manlfeated by the early appearancea

of the "reduotional" grouplnga and la te r development of the more dlaorganlned "aegregational" and "ecattered" figurea.

In thla event the

l a t t e r eould have arlaen a t the atage before the nuclear membrane haa

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80
be#& completely fixed*

In theee eeeee» the ohrcmoeomee, heviag elreedy

etteined th eir metepheee leegth, hut leeklne le o rleetetlo e, elmply
epreed througheet the c e ll mhee the noeleer mcmhrmme fle e lly **hreeks
dome."

Hence the dlsorgenlxed fl^pree mould erlee» not from en eroelon

of the epindle but ee e re e n lt of I te fe llin g to form.

Whether or not

the dleorgenl eetion mould teke the fozm of e "eegregetlonel* grooving
mould then depend either upon eheeee or mhether or not e eufflelemt
emount of Aetl-dlone bed been preeent to here dlerupted the polerlty
of the cell*
4*

In the eeeee of e l l of the ehort t r eetment epperently none of

the e e lle hed been completely fixed end henee "rerereele" mere the gen­
e rel ru le .
In other morde, belem the to x lelty level, Aetl-dlone eppeere to
here e tendency to belt prooeeeee rether then to deetroy el omemte e l­
reedy preeent. I . e . , to be e "fixer" rether then en "In ltleto r" of eetIvltlee*

The teetlng of the effeete of other en tlb lo tle meterlele In

the lig h t of th le hypotheele might yield eone Importent elmee eoneemlng
the modee of eotlone of, e t le e et, eome of theee entl-grovth eubetemeee.

Tr<m the preetleel vlempclnt I t mould e e # thet eneh Informetlon mould
be neceeeery for nelr*"c deelelone eoneemlng the uee of theee eubeteneee
ee beoterlocldee, fungleldee, entl-protozoel poleone, etc.

Ibr exemple,

cognleenee of the feat thet Aetl-dlone, In conoentretlone ebove 8 p|m,
deetroye the mitotic a c tiv ity of the merletemetle tleeuee mould eppeer
to be eeeentlel for detemlnetlone of coneentretlone end methode of
eppllcetlon of the entlblo tle ee e fungicide.

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81

TvæXhmrmarmp I f th# bjpotta#tio#l #n#ly#l# of th# mltô t 1# hr##kdomxi o#n##d by Aotl-dion# d##orlb#d mbor# 1# ###%m#d to b# oorroot,
th#B #om# postulat## #oao#rmlag th# #T#nts In normal mltool# may b#
proposod:
1*

I f th# app#arano# of th# **aff##t#d** prophaa## Indloat##

that Aotl-dlom# eaa produe# thl# s ta llin g #ff#et upon th# amsl#ar
msmbrao# without apparently sta llin g th# d#T#l0]m#nt of th# ehrcmosonas
toward th e ir metaphas# ooadltlon, tmo assumptions may b# made; eith er,
beeaus# of th# slow penetration of th# antibiotio i t has not penetrated
th# nuoleus or Aetl-dlone has a d iffe re n tia l effect upon t hese two
stru ctu res.

Slnee these "affected** prophases occurred regularly in

root tip s continuously treated a f te r a rath er long period of time, the
f i r s t of these assumptions semis le ss probable and a d iffe re n tia l affect
(most lik ely In degree) sssns more acceptable.
2.

la la te r stages of these "affected" prophases (FI. T, F

I t was pointed out th at many (sometimes a l l ) of ths metaphase-appearlng
chromosomes which sessmd to be s t i l l within the nuclear mombrane, were
unorlented.

This permits the conclusion th a t i t may be quite possible

that the orientation of the chromoscaws may be maintained, and possibly
metaphas# alignment directed by the spindle.
5.

I t w ill be recalled that the "reduotional" groupings occurred

in the early hours a fte r the in itia tio n of treatment.

I f Acti-dione i s

considered to behave like a slow fixing agent with a difference In the
degree of a ffec t exerted upon the chromosomes and the other constituent!
of the c e l l , then these "reduotional" groupings give us some evidence
that:

(a) The spindle may be gradually formed; the gap representing the

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at
(b) At l#m#t mcmm part of th# spladl# mmj b# of

und«nr«lop*d

oytoplmmmlo origin fo r. I f d lff# r# n ti# l a o tir lty of Antl-dlon# 1#
mmsimod, then, wh#n th# oytoplamm hm# b##n #ff#ot#d and It# aotlvltl##
r#t#rd#d, th# r«Bk#lnd#r o f th# splndl# eannot form#

Thl# 1# oontr##t#d

with th# vl## of Wad# (140) of th# lntr#nuel#ar origin of aplndl# #abatane##*
4.

Sin##, In th# eomplataly dl#organls#d flgur##, vhioh r# fl# et

th# t o ta l Inhibition of th# aplndl#, th# ehromtld# #T#ntuallj do "fall"
apairt, th# g#a#rally aoo#pt#d ooncopt that th# a p llttin g of th# k#a#tochor# p#r a# la not dapandant npon any aplndl# influonc# la anpportad#
5.

In th# dlaorganlaad anaphaaaa (fjrontiapl### Fig* F, PI* IF,

Fig* G through I) both orlantad and unorlontad ohromatida bar# baan
obaarvad apraad throughout th# call*

I t la raeognlnad that thla d la-

trlb u tlo n w m j hav# ra au lt ad from th a ir "drifting " In th# p a rtially
"fixad" aytoplam»*

On th# othar hand, homavar, thay may b# eonaldarad

aa avldanoa In aupport of th# poaaiblllty th at a t laaat aoma for###
aaaoolatad with chromatid movamant may ba locatad within tha ohromoaomaa thamaalvaa*

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SOMKAKT
1#

Onion root t i p morl#t#m# ##re trontod with th# antibiotio

Aoti-diono in aolntiona of varying eoneaatration# ranging from 1 pim
to 80 p ^*
£•

%# novoTol eoneaatration# wara applied to tha mariatem#

by two main typa# of method#:
(a) A eontinnon# traatmant for 12 howrs*
(b) A ahort traatmant for 15 minuta# a fta r uhioh time tha
root tip# wara tranafarrad to a d iatiU ad water laaoh#
3.

Ineludad for purpoaa# of oompariaon wara two intaxmitta

traatmanta uaing a 40 ppa aolution for 15 minutaa; ia one i t waa
applied ovary hour and in tha othar i t waa applied ovary two houra*
4*

Thara wara thraa typaa of oontrola:
(a) An individual root t i p ooatrol for whioh tha uatraatad
root tip a wara

from a aingla onion a t a a1ngla

time*
(b) Conaaoutiva hour aariaa fo r whioh tha untreated root
tip# wara

a t tha aeama time a# tha traatad

material*
(o) An individual aariaa oontrol fo r whioh tha root tipa
ware taken from tha bulb juat before traatmant waa
initiated#
5#

From amear préparât Iona of tha root tlpa a atudy waa

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64
of tho rarioua typo# of dlTlslon tlg u r m m whieh appearod a t hourly
Interval# fo r a period of twelve hour#.
6.

There wa# a general trend toward a reduction in the number#

of divl#lon figure# following the abort treatment and an eventual eta—
b l lity in theae nimber# in the eomtinuoualy treated root tip # .
7.

With the exeeption of the oytologioally ineffective 1 ppm

abort tree Plant aerie#, the f i r s t fiv e hour period following the in­
itia tio n of treatment wa# found to have been the moat c r it ic a l period
for the obaervatioB of the effect# of Aoti-dione upon meriatematic tiaaue.
8.

Difference# in the re la tiv e frequencie# of the individual stagei

appeared depending upon the concentration and the method of treatment#
9#

On the baaia of the combined data of the f i r at fiv e houra,

differenoea were obaerved in the frequencie# of the aberration# la the
individual atage# with the higheat percentage# occurring in prophaae
and the loweat in anaphaae.
10#

P o aaib ilitiea of the interrelationahip of the eeveral type#

of aberration# a# well a# the cytological and genetioal p o ten tialitie#
of each were diacuaaed.
11.

On the baaia of the hourly v ariation atudy the frequency pereeni

age of the aeveral type# of aberration# appeared to be correlated with tin*
rather than with e ith e r the concentration or the method of tr e a g e n t.
12.

I t waa auggeated that Aoti-dione appeared to have acted in a

manner aimilar to th at of a alow fixing agent upon the dividing cell#;
that is , to h alt proceaaea rather than to in itia te them.

Hie a#aimed

alow fixing action appeared to have manifeated it a e l f as a a ta ilin g of
the noimal change# that take place during mitoaia resulting in the pro­
longation of the duration of each atage#

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85
15.

It

mmeg##t#d that th# degrae of th# fixing action aam

proportional to th# tin# aa ##11 aa th# eonoantratloo and, aa th#
dagr## of th# fixation #aa inar#aa#d, th# a#r«ral typaa of abarrationa
baoaa# apparant.
14.

Sine# no raeorarabl# abarrationa #ar# found, i t naa auggaatad

that the thraahold of to x ie ity may b# very cloaa to that naoaaaary for
the production of th# cytological abarrationa.

Vurthaxmora, tha thraah­

old of each appaara to l i a in aoaia combination of concentration and tin#
batwaan 1 ppm and 5 pgm and batvaan 0 and 15 minataa.
15.

in aararal of tha typaa of aberrant figure# i t appeared that

tha chroBoaoaiaa aamaad to have bean capable of proceeding through th eir
regular morphological changea ahan, a t tha aama time, there waa avidanca
of p a rtia l or t o t a l apindla auppraaaicm.

On thia baaia i t waa auggeated

that A cti^lone may have had a d iffe re n tia l affect upon tha cytoplamic
and nuclear conatituanta of tha c a l l,
16.

Aaauming tha alow fixation action of Acti-diona to be correct,

the following poatulataa concerning a c tiv itia a in normal mitoaia may be
proposed:
(a)

A gradual formation of the apindle.

(b)

Cytoplamic origin of, at laaat, acme parta of the
apindle.

(c)

Poaaibility of tha origin of anaphaae moramanta from
aoma meohaniama ariaing within tha chromatida themaalTea.

17.

Three possible approaches of further inrestigationa are

suggested:

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86
(•)

Th# ###r#h for r#eor#r#bl# #b#rr#tioa# from Aetidlon# should b# oontlnusd by testin g oosiblnstioiis
of eonosntrations b#t###n 1 ppm mxA 5 pjai
bstwosn O aztd 15 mdjmtss*

tlm#

Sueoess in th is sndsavor

would probably ylsld re s u lts of oonslderable praetloal
slgnlfloanoe.
(b)

Tbm hypothesis of the slow fixing action of Aetl-dlone
should be tested using other ehesiioals, particularly
the a n tib io tic s .

Results of such an Investlgmtlon

might lead to a b etter understanding of the mode of
action of anti-growt h substances and could resu lt In
some Important changes In the methods of practical
u tilis a tio n of these m aterials.
(c)

Since the aberrations appeared to have developed during
the early hours following In itia tio n of treatment, i t
appears th at continued observations made during th is
p articular time would yield data from which a b etter
understanding of the fundamental processes of mitosis
could be developed.

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TKXT FIQTOBEB

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

87

KEY
UNTREATED
SAME
HR
UNTREATED
CONSECUTIVE

HRS

15 M I N U T E
TREATMENT
CONTINUOUS
TREATMENT

u n t r e a t e d

T E X T FIG
PER AXIS

I PPM

5 PPM

2 0 PPM

4 0 PPM

8 0 PPM

I A V E R A G E N U M B E R S OF DIVISION F I G U R E S
O B T A I N E D F R O M C O M B I N E D DATA
OF FIRST
5 HOURS.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

CD

■
D
O
Q.
C

g
Q.
■
D
CD

(/)
C/)

"8O

CD

33 ".
CD

AI i tl »1 I I I

CD

■
D
O
Q.
C

a
o
3
T3
O

2 4 # » 10 12
TEXT F it 2
UWTREATEO
CONSECUTIVE HRS

2 4 • B 10 12
TEXT FIG 5
I PPM-19 MIN

2 4 f • 10
TEXT FIG 4
9 PPM-19 MiN

2 4 G • 10
TEXT FIG 9
to PPM-IS MIN

2 4 BB10 12
TEXT FIG 7
80 PPM-19 MIN

2 4 e B 10
TEXT FIG S
40 PPM-19 MIN

IN04VI0UAL
ROOT
CONTROL

*20

*20

*10

*10

CD

Q.
•

T3
CD

(/)
(/)

•10
t p i 1 1 1 1 1 I > 11 I I . g o

2 4 ? 10 12 19
TEXT FIG B
un tr ea ted
SAME hour

TEXT

. . .

. . . . . . . .

.

j qJ

. 1 a 1 . 1 > 1 1 1-1-l- y p l i . l l A l l l A l A t l

2 4 G B I 0
~
2 4 B B K )
"
2 4 G# 10 «
2 4 G B K) *
2 4 GG10 It
TEXT FIG 9
TEXT FIG 10
TEXT FIG II
TEXT FIG It
TEXT FIG IS
I PPM-CONTINUOUS 9 PPM-CONTINUOUS tO PPM-CONTINUOUS 40 PPM-CONTINUOUS GO PPM-CONTINUOUS

FIGS 2 -1 3

HOURLY

CHANGES IN AVERAGE NUMBER

OF DIVISION FIGURES

PER

AXIS

S

09

•o
60
o
X 40

w

“• fO

a. 2 0

12 MR S
T C X T

F i e

1 4

T E

U N T R E A T E O

j-eo
4
*-

O N E

P P M

F O R

15

M < N U T f S

* «O

4 0

w

tO-

T C X T

F i e

I B

F I V E

F P M

F O R

1 5

M

O

20

I N U T E S

T E X T

F i e

17

T W

E N T Y

P P M

F O R

I S

M I N U T E S

K
M

I C O

B

IS

BO

*

o
oc
w
a.

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6 0

B O

w

K T

­

B O

BO

*40^
20

5
T E X T

F i e

I B

B

F O R T

V

M R S
P P M

K E Y

T E X T

F O R

P R O P M A S E

—

—

( P E R C E N T A G E S

TEXT

FIGS

14-19

HOURLY

—

m

e

t a

p

h

a

C O N V E R T E O

VARIATION

s e

—
T O

IN

F I G

• —
B A S E

1 9

E I G H T Y

P O S T
O F

-

M E

P P M

T A P M A

F O R

I S

M l N

J

T E

S E

5 0 )

FREQUENCIES

STAGES

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OF

INDIVIDUAL

?

90
#0
KCV

•O

m
m
oPM
A
SE
- —

-

—

w

ME TAPMASE

40

PO ST -M E T A PM A SE

w

HRt
TE*T
It0f>

itoH

lOO

lOO

w

SO

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FiQ t o

ONE

P P M - CONTINUOUS

•o -

w

W 40

K 40

to

to

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FIO t l

FIVE

PPM - C O N TIN U O U S

TEXT

looh

isoh

140-

140-

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TEXT FIG 71 'SCATTERED' MCTAPHASES

^

SHORT TREATMENT ^

PERCENTAGES OF A B ER RA NT

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96
8
KEY
UNTREATED
SHORT

TREATMENT

CONTINUOUS

o
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T E X T FIG. 7 3 . M E A N P E R C E N T A G E S
OF P R O P H A S E a M E T A P H A S E C E L L S
C O N T A I N I N G " S E G R E G A T I O N A L " OR
" R E D U C T I O N A L " G R O U P I N G S F OUND
DURI NG F I R S T F I V E H O U R S

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

FIATES

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

FIATS I
•SGRUAIXT" ORGANiaD) FIOCRZS

TiaORE
A.

Propham# frcm root tip takoo alght hoars a fto r in itia tio a
of short traatment with 1 pgm solution*

B*

ProsMtaphass from root tip taksn a fto r two hours of eon—
tinnoos troatSMnt with 20 pjm solution*

C*

Motaphaso from untroatod material*

D*

Early anai*aee from untreated material*

E*

Middle anaphase from root t i p taken one hour a fte r begin­
ning of short treatment with 1 pjai solution*

F*

Late anaphase from untreated material*

O*

Early telophase from root ti p taken a fte r one hour of con­
tinuous treahaent with 5 ppm solution*

H*

Later telophase from root t ip taken one hour a fte r begin­
ning of short treatment with 1 ppm solution*

I*

Late telophase accompanied by two in te r phase c e lls from
untreated material*

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

97

FUVTK I

'A
.

#

I

# a
Him
y

#

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

#

ê

I

FIAOX II
A SERIES OF "REDOSTIGNAL** GR0DPINQ6
FIGRXOE

A.

FiropbaM "r#duetlomal" grouping from root tip tnkon n fto r
MM hour of oontiimoun trontmont with 40 ppw nolutiOB.

B«

Pronotnphnso "roduetionnl* grouping from untrontod mntorinl,

0.

S p lit

motnphnn# fromuntrontod mmtorinl.

D.

S plit motnphnn# from

root tip tmknn a fte r two hours of eon—

tinuoue treatment with 40 p%m eolntion*
E.

S p lit anaphaae from root tip taken a fte r two and one-hailf
hours of eontinuoos treatmsnt with 40 ppm solution*

F*

S p lit

anaphase from untreated m aterial.

O.

S p lit

telophase fromroot tip taken a fte r

six hoursof

eontinuous treats&ent with 20 ppm solution*
H*

S p lit telophase from untreated material*

I*

S p lit

telophase fromroot t i p taken three

hours a f te r

beginning of short treatment mrith 5 ppm solution*

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

98
PLATE I I

f

^

^

A

#

••
%b

•

/
E

#

k

■ 'é

f

I

•» I
*

4

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I

FIATS I I I

A SERIES or «SBGREGATIQNAL” GROÜPIKOS
FIGURE
A*

Metaphm## "##gr#g#tlom" ImrolTinE equal ohroaoaoaie
mmberm from root tip taken a fte r four hours of
eontinuous treatment with 80 pgm solution.

B.

Two adjacent metaphase c e lls, one (upper) imrolvlng
squal, the other unequal chromosome nisabers, from
root t i p

a fte r four hours of continuous

treatment with 00 pgm solution.
C.

Metaphase " ssEregatiom" iarolvlng equal chros&osome
numbers from root tip taken a fte r four hours of
continuous treatment with 80 ppm solution.

O.

Unequal netaphase "segregation" in which chromatids
have completely "fallen" apart from root tip taken
a fte r three and one*half hours of continuous tre a t­
ment with 40 pxm solution.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

99
PIATK III

A

iT C '

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

FIATS 1 7
A SERIES 07 "SCATTSRSD" 7IG0RS8 07 THE DISORGANIZED TXFS
nOORX
A.

Dl#oremmlz#d prcmetapha#e fr o m root t i p tmk#n a fto r aix
hours of ooBtinuoua trea^aent with 80 p%m aolutioa*

B •» C •» D*
"Seattarad* mataphaaaa frcm root tlpa takaa a f ta r four
hours of aontimious traatmamt with 80 ppm solution»
%»

Normally orcanlsad mataphasa shoulng aridaaoas of apindla
supprassioa in tha typa of kinatoehora s p littin g , from
root tip takan a fta r four hours of oontinuoos traatmamt
in 80 ppm solution»

7»

"Seattarad" mataphasa in which chromatids hara "fallan"
apart, from root tip takan six hours a fta r the beginning
of tha short traatmant with 80 ppm solution»

0»

Disorganized anaphase inrolring oriantad ahromatids taken
from a root tip a fta r four hours of eontinuous traatmant
with a 1 ppm solution»

^

H»

Disorganised anaphase inrolTing both oriantad and unoriantad
chromatids taken from root tip six hours a fta r the in­
itia tio n of the short traatmant with a 40 ppm solution»

I»

Disorganised anaphase inrolring mostly unoriantad ahromatids
takan from root tip a fte r four hours of continuous tr a a t­
mant with 80 ppm solution#

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

100
PLATE IV

'A ,

<

4

»
V‘ "

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

T*

FLATS V
A SSRIS8 OF CSLIS SHOWING INDICATIONS OF TOXIGITT

FIGCRS
A.

D#t#rlor#t#d iat#rpham## from root ti p takoa oftor oloren
hour# of eo&tlaixmo trootownt «1th SO p%m oolotion*

B#

"Affoetod" promotopbmoo from root tip tmkon of to r four hour#
of ooBtiBuono troo^koBt with 80 ppm solution*

0.

"Affoetod” propbooo from root tip tokon of to r fivo hours of
oontinuoos trootmont with 80 pgm solution*

D*

"RoTorting" prophsso f r om root t i p tokon fivo hours o fto r
initlotiom of short trootmont*

B*

"Noorotio" propbooo from root tip tokon oftor t hroo hours
of oontinuous trootmont « ith 1 ppm solntiom*

F*

"Clumpod" motopboso from root t i p tokon oftor six hours of
oontinuoos trootmont with 80 p%m solution*

G.

Lots snophoss with

ehromosomos frcm root tip tokon

oftor ton hours of eontinuous troaUoont with 80 ppm
solution*
H*

"Affootod" tolopbooo with chromosomo "bridgos" from root tip
tokon oftor throo hours of eontinuous trootmont with 40
ppm solution*

I*

Tolopboso with unoquol soporotion of ehrcmotids from root
tip tokon oftor ni no hours of eontinuous trootmont with
40 ppm solution*

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

101
PLATS V

4
I
%
»

€
#

#
i

a
y

*#
»

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

\

PIATZ VI
laSCXLXiüSBOOS
naoRB
A.

Ammphm## of m polyploid dlTioloa frcm root tip tokon thro#

hour# of tor inltlotioA of ohort trootmont with 1 ppo
eolation#
B#

Metophoeo with ohromoeome frogpwnt from root ti p token one
hour of te r in ltlo tlo n of ohort treotment with 40 p%
m
eolation#

0.

Binooleote e o ll from root t ip token of te r five hour# of
eontinuooe treotment with £0 ppm eolation#

D#

IVinooleote o e ll from root t i p token of te r twonty-one
houre of eontinaoue treotment with 80 ppm eolation#

B#

Rodooed oell from root tip token of te r two houre of con*
tlnaoue treotment with £0 ppm eolation#

f.

Rodooed o e ll from root tip token of te r eiz houre of eon—
tinuoue treotment with 40 ppm eolation#

O#

Dieorgonloed telophoeo from root tip token four houre
of te r in ltlo tlo n of ohort treotment with 9 ppm eolation#

H#

"Droneitionol" metophoeo from root tip token of to r throe
houre of eontinmoae treotment with 80 ppm eolation#

I#

Unipolor onophoee from root tip token of te r two houre of
eontinaoue treatment with £0 pjm eolation#

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

102
PTATK TL

#

I

é \

• ' ft

D *

a
•

V

«

a

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

c

BXBLIOGRAFHT
(1)

(2)

Allen. N. S. (1949)
lui. S. Tbesls: Induoed mltotio "reduetional grouping#" in root
tip# of tetraploid te ^ e s e n n tia .
UniTeraltj of Wlaeonsin.
.
a« B. Wllaon and s. Powell (1950)
CcwparatlTe effaota of oolebiolne and aodlun nuoleate on aomatio
ohroaoaoaiaa of Allium and Tradeaeantla.
Jour. Bared. 41:159-163.

(3)

Angulo Carplo, H. Dolorea (1949)
Sfaetoa o-«aitotloom da la morfina.
Genetlea Iberioa 1:41-47.

(4)

Arnaaon, T. J . , X. Ounmlng and J . Splnka (1948)
ChrcBoaome breakage in plant# Induead by radloaetlve ptaoaphorua (P^)
Soienea 107:198-199.

(5)

Auerbaoh» 0. (1950)
Dlffaraneea between effeota of ehemloal and ptayaieal nutlgana.
Pub. dalla Sta. Zool. dl Bapoll 22 (auppl. ) :1-19.

(6)

Barber, H. N. and H. O. Callan (1943)
The affecta of cold and colchicine on mltoaia In the newt.
Proc. R. Soo. B. 151:2M-271.

(7)

Battaglia, m illo (1949)
Sull'aalone d e ll’etil-oarbamato (uretano) e del clcloeall-aarbeatato.
Caryologia 1:229-247.

(a)

_________ , (1950)
Nuore aoatanse Indueentl franmentaslone oromoacmloa.
Pub. della Sta. Zool. dl Napoli 22:125-157.

(9)

___________ , (1950)
Oaserraalonl aull*aalone oltologloa dl aloune aoalanze coloramtl.
Caryologia 2:223-228.

(10)

Bauch, Robert (1948)
Trypaflavln ale Typus dar Chrcnoaoaengifte.
Naturirlaa 34:346-347.

(11)

Beal, J . M. (1950)
Chroa&oacae aberration# In onion root# frcm plant# gromi tn #»
atmoaphere containing 0^*0g.
Amer. Jour. Bot. 37;660.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

104
(12)

Bemmm, H* W. and R. L. Ring (1996)
Ab «xpariaantal r%udj on mltoal# in tha aomatio oalla of ahaat.
Biol. Bull. 75:189-007.

(13)

Borgar, G. A.
(1944)
Xzparimantal atudiaa on tha cytology of AUiim.
Torraya 44:41.

(14)

,

and S. R. Witkna (1943)
A oyto logical study of C-4litosis in thapolyaomatio
plant
Soi nan ia Olaraoaa. with oomparatiTo oboorratioas on A)] i f
Oana.
Bull. Torray Bot. 70:457-466.

(15)

,
and X. R. Witkua (1946)
Polyploid mltoaia aa a normally ooourring factor in tha davalopmant of A llim Cana L.
>. Jour. Hot. 33:785-787.

(16)

and X. R. Witkua (1948)
Oytologioal offacta of Alpha-naphthalono acetic acid.
Jour. Barad. 39:117-120.

(17)

Bishop, Ohaa. (1950)
D iffaraatial X-ray aanaitivity of ^radaacantia chromoaomaa
during tha mitotic eye la .
Qanatica 55:175-187.

(16)

Blakaalaa A. P. and A. G. Arary (1937)
Methods of inducing doubling of chromoaomaa in plants by
traatmant with colchicina.
Jour. Barad. 88:393-411.

(19)

Brown, M. S. (1947)
A case of apontanaoua reduction of chromosome nombar in somatic
tissue of cotton.
Anar. Jour. Bot. 34:364-388.

(20)

Bruaa, A. and A. Cohan (1936)
Xffacta of Colchicina and related aubatancca on c a ll division.
Biocham. Jour. 30:1363-1368.

(21)

Chargmff, X., R. M. Stewart and B. M
agcaanlk (1948)
Inhibition of Mitotic Poisoning by mcao-inoaitol.
Scianca 108:556.

(22)

C hristoff, M. and M. A. Christoff (1948)
Maioaia in the aoamtie tiaaua rcaponaibla for the raduction of
chromosome nimbcr in tha progeny of Bicrocita: Boppaanom Schuit.
Oanatica 33:36-42.

Reproduced with permission o f the copyright owner. Further reproduction prohibited without permission.

10#
(S3)

Ohmrf, J . «ad VI. Slooka (1948)
VliT broBO M
t ko^ano^h Troholaeh A lllta Caw.
Splay Lakai^aka Vkk. Maaarykory DbIt . (Brno) Sj^l-lS.

(84)

Coraaan, I . (1947)
Tha raaponaaa of onion and I lly mltoaia to eowarin and
paraaorbio aold.
Jour. Szp. Biol. 83:898.

(85)

D*Amato, V. (1947)
AgglutInanloma aroamtlca ("atiekinaaa**) In A lllw Cana a
a ltr a Lillaaaaa,
A ttl Aoaad. Lineal (Roma) Cl. Sol. Fla. Matl. Nat. Sar. 8A
3:135-140.

(86)

________. (1948)
Contrlbttto a l problama della agglutlnasiona oromoacmloa
(atloklneaa) apontanaa.
A ttl. Aooad. Linoal (Roma) Rand. 0. Sol. Fla. Matl. Nat.
Sar. 8A. 4:94-98.

(87)

(1948)

Tbm off act of oolohlolna and athylana glyool on atloky
ohromoacmaa in A U lw Cana.
Haradltaa 34:63-105.
(88)

________, (1948)
Htttaaloni oromoaomloha apontanaa In Nothoaoordma fragraaa.
Knnth.
Caryologia 1:107-108.

(89)_
__________, (1948)
Rloharoba a u ll^ a ttlr lta oltologloa dl alounl oompoatl organic1
oon partloolara rlgoardo a lia oolohlo Ino-mltoal a agll a ffa tl
toaalol.
Caryologia
49-78.
(30)

________, (1949)
Tha affaot of m-lnoaltol on C-mltoala and C—
tomor reaction.
Caryologia 1:358-361.

(31)

_________, (1949)
Praprophaaa Inhibition of mltoaia In root marlatoma.
Caryologia jL: 109-181.

(38)

, (1949)
^ ^
S ull^attlT ita nntagana a aul tlpo dl mutaalonl oromoaomloha
Indotta dall'aoanaftana.
Caryologia 1:801-806.

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106
(35)

(34)

(55)

U'jWmto,
(1949)
9nll*iapl«80 4#1 ffUMMno ocm# ag#Dt# poIiploidimBut*.
Caryologia
809-322.
(1950)
iSeehpeeoseee brooking a e tir lty of chomiealo a# atudiod by
the A llim C
Gena
OPO teat*
Pub* della Sta*
StaT Zool* di Napoli t t (auppl*) 156-170.
_______(1950)
Studio atateatleo d e l l 'a t t i r i t à mutageua dell* aeridina e
derirati*
Caryologia 3:339-297*

(56)

and M
* O* Aranni (1949)
Aplmo oontrib u te a lia oonoaeensa d e ll* a ttiv lta oitologiea e
fiaiolegiea delle aoatanse di ereaoita aul te a t a i i j m Cena L*
caryologia 3:51-54.

(57)

______
and K* O* Aransi (1949)
studio oomparativo dell*attivit& oitologiea di aloune eseense*
Caryologia 1:175-193.

(38)

. and A* Guatafaaon (1948)
s iu iie a on the ezperimental control of the mutation prooeaa*
Heredltaa 34:181-193*

(39)

Uarlington, 0* D* and y. C. Koller (1947)
The ahemioal breakage of ohromoaomea.
Heredity 1*187.

(40)

Dermaa, H* (1940)
Colohioine polyploidy and technique*
Bot. Rev* 6:599-635.

(41)

(1941)
intranuclear polyploidy in bean induoed by naphthalene acetic aold.
Jour* Hered* 33:133-138.

(43)

Deyaaon» Guy (1948)
Sur 1*effect tropoelnetlaant dec agente a mltoclaaiquea.
Bull. Soo* Bot, (Rranoe) 95:306-311.

(43)

Bdeard, Pierre (1949)
Action du bomeal aur la division oellulaire ehez l*Alliisn Cepa L.
Bull. Soo. Bot. Trance 96:330-333.

(44)

Xhrenbergf 1** A. Ouataffson, A* Levan and U. von Wsttstsin (1949)
Radlophoa^oroua, aeedllng le th a lity and ohrcmoeome disturbances.
Heredltaa 35:469-489.

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107
(45)

Sigati, 0. J . (1947)
Oolohloina Bibliography#
U oardlo (Oinolnnotl) 10%65-114#

(46)

Srlokoon» R# O# (1948)
Ppotooneeonln am a mltotio Inhibitor#
Solmnoo 108;533#

(47)

Fmroraky, H# 7# (1939)
M
om polyplol&y—
IndooIng ohmmiloalm.
C. R# Aoad# U#R#S#S# 25:71-74#

(48)

Parguoom# J . (1939)
The ueo of ohamleal potential# aa Indloea of toxloltj#
Proo# R. Soo# London Ser# B# 127:387-404#

(49)

Ford» J# H# and B# B# Leaoh (1948)
lotl-dlame, an entiblotlo from Streptcmaroaa arlaeua.
Jour# Aner# Oham# Soo# 70:1223-1226.

(50)

I^ralK-Lelmald» J# A# (1948)
Polyploldle b lj planton door ohaail aohe varblndingen#
Chrom. Bat. 104:321-334#

(51)

Frlaaner# Ray C# (1920)
Dolly rhythm# of elongation and oell dlrlalon In certain root##
Amor# Jour# Bot# 7:380-407.

(52)

Oallnoky, I# (1948)
The o ff act of aartaln phoepbatea on mltoaia#
Genetloa 33:610-611#

(53)

(1949)
The o ff act of certain phoaphatea on Mltoaia In AUlmm roota#
Jour# Hared# 40:289-295#

(54)

Oentoheff, G# and A# Onetafaeon (1939)
The double ohromeoome raproduotlon In Splnaola and Ita oauaaa#
XI# An x-^ray experiment#
Heredltaa 25:371-386.

(55)

Glbaon, P# B#, R# A# Brink and M# A# S t a f aim (1950)
The mutagenic action of muatard gaa on Zee mayo.
Jour# Heredity
232-238.

(56)

Glooe# A. C# (1947)
Radiation# and o e ll dlTialoma#
quart# Rev# Biol# 28:283-282#

(57)

Gllaa, B# H# and A# D# Conger (1950)
Chrcmoaoma Inter#hangaa induced in Tradeaeantla mloroaporaa oy

faat nautrona from uranium flaalon#
Jour# Call and Comp# Phyaiol# 35 (auppl) 83-88#

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106
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OulAoehat» M. (1940)
S«r
a o d iflo « tlo u &## r##otlonm pbymloo-ohlmlqu## d#
la oa lla i# v#e#tal# proreqo### par la# aubatano## aitolnhibitrle##.
0. R. À#ad. Sol. SIC! 579-960.

(99)

Ouatafmaan, A* (1999)
Tha ia ta rra la tio a of Maioaia and Mitoaia.
A#amoapa%*y.
Baraditaa 25:989-922.

(90)

(91)

Tha maobanimm of

,andJ . UaàEmj (1948)
tka canatioal affaota of moatard gaa aubatainoaa and nautrona.
Haraditaa 54:371-989.
Hoaklna» 0. L. (1954)
AjMMlova aagragation of a trip lo id tonato*
Jour. Haradity 29:221-224.

(92)

(1947)
Tha auhdiTiaioB of tha ahromoaomaa and th air multiplioation in
mon dividing tiaauaa: Poaaibla intarpratationa in tama of gana
atruotura and gana action.
Amar. Bat. 81:401-434.

(95)

(1948)
SagragatloB and raduction in aomatio tiaauaa.
tiona in Allimn oapa.
Jour. BaraA. 3?:511-925.

I# I n itia l obaarva-

(94)

. and K. C. Ohong (1950)
BagragatioB and Raduction in Somatic Tiaauaa. IT. Raductlonal
grottpinga induead in Allim: Capa by lou tomparatur a a.
Jour. Barad. 91:15-18.

(95)

. and L. Chouinard (1950)
fliomatic raduction: Diploid and trip lo id roota and a diploid
ahoet from a tatraploid Rhao.
Oanatica 55:115.

(66)

KaUioott, W. 1. (1904)
Tha d aily periodicity of call-diviaion and of elongation in the
root of
Buli. Torray not. Club 91:529-590.

(67)

Rihlnan, B. (1950)
8—
Bthoryaaffain, an ideal induoar of atructural chrcmoaoma changea
in tha root tipa of A^iun capa.
Xcptl. C all. Raa. ^:T35—
d8.

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10#
(68)

«ttd
(1949)
th# eytolesieal eff#ot of ooffolm.
Béroditao 38%109-111.

(69)

Kodoai» M. (1948)
S od iv rl#om# BQOloato and n lto o ia.
Indnotion of morghologloal
ohango# la tha ohromoaom## and of ahnormalltl## in a ito tio
dl violon# in the root nerlotomo.
Jour. Harod. 39%387-535.

(70)

Kootoff, if. (1938)
Oolohlolna and aoanaphthana ao polyploldloln# aganto.
Nature 142:785.

(n )

^
. (1938)
Irregular Mitool# and Maloolo Induoad by Aeanaphthana.
Nature 141:1144-1148.

(72)

. (1948)
Atypical growth, abaoxmal mltooio, polyploidy and ohrcamooaaa
fragmentation induead by haznehlorooyolohaxana.
Nature 162:845-846.

(73)

Leaoh, B. 2 ., J . h. ford and A. Vhiffen (1947)
Aoti-dione, an antibiotio from atrautcouroao orloeuo.
Jour* Amer. Uhom. aoo. 69:474-475.

(74)

Loferre, J . (1939)
similitude deo aotiomo oytologiqueo ezeroiao par la
ph4nylurdthana at la colohioina our leo plantuleo Wgétalao.
C. R. Aoad. 30. 208:301-304.

(75)

Laite-Rio, L. F. (1949)
Sotudoe da oitologia azparimantal. I . Contraoeao doo
eromoocoMO paloo oatloeo.
Bol. 3oo. Portugueoa Giono Nat. 17:111-138.

(76)

Levan, A. (1958)
TIm effaot of colohioina on root mitooio in A llli» .
Bereditao 24:471-486.

(77)

______, (1939)
(Sytologioal phenmmena oonnaotad with root owalllng oauead by
growth oubotaneao.
Haraditaa 88:87-96.

(78)

_____, (1939)
Tatraploidy and Octoploldy induoad by colohioina in Diploid
petunia.
Bereditao 28:109-131.

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110
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A. (1940)
Th# effect of eeenephthene end eolohiolne on nitoeie of
A l l l t i ead O o le h ie i»
Hereditae 24:262-076.

(80)

. (1944)
X xperlaeatallj ladueed ehlorophyll mteAte in flex.
Bereditee 30:220-230.

(81)

. (1944)
Votee on the ejrtoloor of Dloeedl end Bellevelle.
Bereditee 30:216-224.

(82)

.

(1945)

dytologioel reeetione indooed by inorgenie eelt eolutione.
Hetnre 186:701-702.
(83)

. (1947)
Stndiee on the eenphor reeetion of yeeet.
Bereditee
407-014.

(84)

. (1949)
R e inflneBoe on ehronoeonee end mitoeie of ohemieele, ee
etndied by the Allien t e s t .
Proo. X l^ th Gong# of Oen. (Bereditee# 8nppl. Vol.): 320-037.

(85)

(86)

______, end T. Lotfy, (1949)
Mephthel one eoetie eoid in the A llim te e t.
Bereditee 35:337-073.
. end O. Oetergren (1945)
The meehenlm of emltotio eetion.
Bereditee 29:301-443.

(87)________# end J . B. Tjio (1948)
Ohrcmoeome frem ontetioa indneed by phenol# #
Bereditee 54:200-251.
(88)

. end J . B. Tjio (1948)

ïaSuetion of ohrcmoeome fregmentetion by phenol#.
Bereditee 54:405-484.
(89)

Lewie# A. C. (1901)
Oontrlbutione to the knowledge of the phyeiology of Keryokineeie
Bet. Gee. 32:424-426.

(90)

Loreneo Andrew# A. (1901)
Aeoion de Verte# Selee Aloelinee Sobre le Dirteion Celuler
on A llim Cepe.
de AalêT7l74-I86.

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I ll
(91)

Lovel###,
«Bd 8. R#v#U (1949)
H«« #vld#B«# OB th# mod# of «otioB of "mltotio polooBo."
Notvr# 164:938-944.

(92)

Lodford, R. J . (1936)
Th# «otioB of tozie #ub#t«Bo## opon th# dirloloB of normal
aad mallgoant o#U# la v itro aad vivo.
Aroh. f . #Kp* Z#Uf. 18:411-441.

(93)

Maofhrlan#, X. W., A. M. Mooolog, aad M. H. Ryan (1951)
Xffoot# of motor oouro# oa toodoity of moreurlal poiooao.
I . StaadardlsatlOB of prooodnr# la toot# aolng A llim roota.
Jour. Horod. 48:98-99.

(94)

Mohro, P. M. (1949)
Xffoot of oalfaallmaldo oa mltotio dlvlalon of pollon gralaa
of Xphodra.
Bot. Qaa. I l l ; 98-63.

(95)

Moyor, Jam## R. (1938)
Modlfloatloa of mltoaia by ehomleala.
Soloaoo 108:188.

(96)

M lttlor, Sldaoy (1990)
OiaruptloB of mltool# la oaloa root oollo by thyroid pomdor.
OoBOtlo# 39:124-189.

(97)

U lttlo r, Sldaoy aad R. H. Harmaa (1990)
Dlaraptloa of mltoaia by doaloatod thyroid tlaauo.
Soloaoo 111;548-590.

(98)

Nobol* B. R. (1937)
Oytologioal obaorratloaa oa eolehlolao.
Biol. B all. 73:35W98.

(99)

(100)

(101)

(108)

, (1938)
Oolohlolna and Aoonaphthono aa Polyploldlslng agoata.
Nataro 148:897.
, and R. L. Ruttlo (1938)
l!ho oytologioal and gonotloal algalfloaaoo of oolohlolna.
Jour. Harod. 89:5-9.
N&aao, B. (1904)
Uebor dlo Xlamlrkang doa Chloralhydrata auf dlo Komund
^ U th o llu n g .
Jahrb. f . mlaa. Bot. 39:649-730.
, (1986)
^
EuTtl polar# t o i lungafIguron und vogotatlv# ohromoaomonroduktloa.
Blologla Ooaoralla 8:96-103.

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US
(103)

Mcvoomt» Smrl H. #md R . B. Wallae# (1949)
Obreeioeeeal and aaslaar aWrrmtlon# iadoBad by altraaoml#
vibration#.
Amar# Jour. Bot. 38:250-236.

(104)

Nlobol#» C. (1941)
Spoataaoooa ataromoaono aborratloa# la AUlma.
Gaaatla# 26:89—
100.

(105)

Novlok, A. and A. H. Sparroo (1949)
Tha affaota af nltragan muatard on mltoala la onIon root tlp a .
Jour. Harad. 40:13-17.

(106)

Nybom M. aad B. Knutaaon (1947)
InvaatIgatlona on o-a&ltoala la AUlam Oapa.
Baraditaa 33:820-233.

(107)

uatargraa, 7. (1944)
Aa a ffio la a t ohaaieal fo r tha lnduotl<m of atloky ohromoaomaa.
Baraditaa 50:213-216.

( 106 )

___________, (1944)
Colablolna mltoala, ohromoaoma ooatraction, marooala and protaln
ohaln folding.
Baraditaa 30:429-467.

(109)

, (1950)
Oanaldaratlama on aema alamantary faaturaa of mltoala.
Baraditaa 36:1—
16.

(110)

., (1950)
Cytologlcal atandarda for tha quantltativa aatlmatioa of aplndla
dlaturbanoaa.
Baraditaa M :371-382.

(111)

. and A. Lavan (1943)
Tha aonnaatlom batmaan o«ailtotlc a c tiv ity and matar aolublllty
In aoma momaoyalla oompounda.
Baraditaa 29:496-498.

(U2)

Patton, R. L., and B. R. Nabal (1940)
rrallmlnary obaarvatloaa oa phyalolagloal and oytologioal affaota
af aartaln hydraaarbona on plant tiaauaa.
Amar. Jour. Bot. 27:609-613.

(113)

Patau, K. (1950)
A carralatlon bat aaen aaparatioa of tha tmo ohroou>aoma groupa
In acmatla raduction and th a lr dagraa of homologoua aagragatloa.
Oanatloa 35:128.

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113
(114)

PoUtM r, 0« (1931)
Dl# Z#Ut#llimc ##tar#ad und anoh d#r Hnrkos#. Xia B#ltr#c
sur Kanntni# d#r 8torun##n d#a Xarat# 1lungrhythmu#.
Zaltmehr. f . Zallformh. U : 334-363*

(119)

Quldat# P. and H. H itlar (1946)
Obtantion da planta# polyplolda# ar traltamant \
1*haraahlaroojralohatana at au sulfura da polyohlorooyolana*
Ccmpt# Rand, Aoad, Sol* 238:833-635,

(116)

Ramdolf, L, 7, (1990)
Cjtoloelaal and phanotjrpioal affaota Induoad in Main# by ra js
and tha Bikini Tant Abla atomic bomb.
Jour, Call, and Comp, Phjsiol, 35 (suppl, 1) 103-117,

(117)

Roaanfiald, M, (1938)
Tha nation of athar on c a lls in m itosis,
Arab, axp, Z a llf, Bd, 18,

(118)

Bosanfiald, M, (1933)
Xxparimantal modification of mitosis by ammonia.
Arch, azp, Z a llf, Bd, 14,

(119)

Satina, S., A. ? , Blakaslaa and A. Arary (1937)
Balancad aad unbalancad haploids in Datura.
Jour, Harad, 88:193-808,

(120)

Sax, Karl (1990)
Tha affaot of x-rays on ohromoaoma structura.
Jour, Call and Comp, Phyaiol, 35 (suppl, 1) 71-61,

(121)

Shiganaga, H, (1937)
An axparimantal study of tha abnormal nuolaar and c a ll
dirisio n s in lirimg c a lls ,
Oytologia, 7DJII Jub, Vol. 1:484-478,

(122)

,(1949)
Xxparimantal studias of abnormal nuolaar and c a ll dirislons,
IT, Obsarrations mith lir in g c a lls of tha affaats of high and
lam taaparaturas,
Cytologia 14:122-134,

(123)

Shimosmra, T, (1939)
Oytologioal studias of polyploidy induoad by colchicina,
Cytologia
488-494,

(124)

Shmuck, A, (1938)
,
_
Tha ohsminil nature of substancas inducing polyploidy in pianxs,
C, R, Acad, Soi, UJl.S.S, 19:189-192,

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114
(129)

(126)

(129)

(128)

(129)

ShBWkÿ
«Bd A. Ottseev» (1989)
Oh«ml#&l Btroetw# af oubetona## grodaelm* poljploldj la ploBt»,
G#R* Aaod • Sol* G*R*3*S# 24*441^444•
_
_
A. OuBsara and O, I l j l n (1939)
ik# eorranpondanog b#t###m tha affaota of oaamaj^thana in
planta and tha oarolnoeanla aomponnda npon animal tiaaua.
Bioohimija 4;490-421.
Simonat , M. and H. Quinaohat» (1939)
Oompoxmiaon da Inaction aur la Bld at la Lin da divaraaa
oubataneaa provoquant daa onomaliaa da la earjoeindaa.
C. R. Aoad. So. 208:1669-1669.
and M. Ouinoohat (1939)
t*aotion da qualquaa ddrivda halocdnda a t nitrda daa oorburaa
ojeliquaa aur laa nitoaaa dana loa eallulaa végdtalaa.
C. R. Soo. Biol. 132:455-459.
Sinka, Bomio (1941)
Tha hydration and dahydration phanomana in mltoaia.
Apparont ohangaa in atruotura of tha nuolaua.
Cytologia 12:1-13.

III.

(130)

Solomon, M. D. and J . A. %pont (1941)
Praliminary raport of Poriodioity and Rhythodoity of mitotio
phaaaa of root tlpa umdor varying light oonditiono.
Tkona. Kan. Aoad. Soi. 44:202-209.

(131)

Stalnaggor, 1. and A. Lovan (1949)
Comatitution and o mitotio ootivity of lao-oolohioina.
Baraditaa 53:585-396.

(132)

. and A. Lovon (1949)
4ka oytologiaal affaot of ohloroform and oolohlolna on AUimm.
Baraditaa 33:515-525.

(133)

. and A. Lovun (1948)
Tha oHoitotio qualitlaa of oolohioina, trimathyl oolohloinio
ooid and tvo phonanthrang dorivativoa.
Baroditaa 34:192-203.

(134)

Ikaub, H. P. (1941)
Sffoot of aulfanllomida and othar aulfa oompounda on nuolaar
oomditioB in planta.
Jour. Earad. 32:197—
159.

(135)

Vaaroma, A. (1947)
Bnparimantal atudiaa on tha influanoo of DDT inoaotloido upon
plant mltoala.
Haraditaa 33:191-219.

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US
(136)

Vmmderlya, L#om (1948)
ScMtie mitomi# in th# root tip of All!mm C#g# - A r#vl##
• bA R#orl#&tatlon«
Bot. R#t . 14:870-318.

(137)

V##ghh, J . R., J . L. Loekaood, O* S. R#nd#r# and C. B#ma#r (1949)
Th# aotion of Aoti-dion# on plant tl##n# and upon oartain fnngl.
Mioh. A ^ . Ixp. (iuart. Bull. 31:456-464.

(138)

TlUara, R. (1940)
Xtuda oytologlqu# d# I'a e tio n du thymol aur laa eallaa rdgdtalaa.
C. R. Boo. Biol. 133:806-206.

(139)

Wada, B. (1940)
Labandhaobaohtungum ubar dia Blmrlrkung of Colohioina auf dia
Ritoaa, inabaaondara ùbar dia FTaga Spindalfigur.
Cytologia 11:93-116.

(140)

. (1980)
K a maohanimm of mitoaia baaad on atudiaa of tha aubmioroaeopio
atruotura of tha living atata of Tipadaaoantia c a ll.
Cytologia 16:1-86.

(141)

nrhiffan. A., N. Bohonaa. and R. L. Baaraon (1947)
Tha produotion of an antifuagal antibiotio by Straptomyoao
ariaaua.
Tour. Baot. 58:610-611.

(148)

Wilaon. O. B. (1950)
Oytologioal affaata of aoma antibiotioa.
Tour. Harad. 41:886-851.

(143)

(144)

and K. 0. Chang (1949)
Sagragation and raduotion in aomatio tiaauaa.
Tour. Harad. 40:3—
6.
Witlcua, X. R. and C. A. Bargar (1944)
Varatrina, a nam polyploidy indueing agant.
Tour. Harad. 35:189-133.

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TABLï I I I

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REPORTE) RESULTS CF APPLICATION OF CERTAIN RIYSICAL STIlïUU TO KERIST&ATIC TISSUE

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0
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STDCDIDS

k A T R IA L

M IT O n C Clrc iE
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S.

ATOËIC RADIATIONS

M AIZE SBtD

FAST NEUTRONS FRŒ
ÜR A H n» F IS S IO N

TRADESCANTU
MICROSPORSS

t

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—

REFERENCE

B CLO

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NEUTRONS P R d i CYCLOTRONS DORMANT SEEDS
OF BARLEY
TKHSRATURS ( lo « ) mod
SODIUN NDCIEATS

■
D

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TDŒEKATURE ( lo v )

TRADE9CANTIA

im PERATURE (lo w )

A lX Itk CEPA

üLTRACSmRIFTJUlNO

O m K IN A T n »
«BEAT SEED

ULTRA SONIC VIBRATIONS

122

X
X

64

X

X

X

X

12

NARCISSOS

X

103

X-RAYS

A IL IU M CEPA

X X

X-RAXS

FLAX-SEEDS

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S F IK A C U ZEA
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APPENDIX n

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PERCamiŒS 0? TOTAL DIVISION nCRKES OF IKDIVIDUAL STAGES
A. SHORT IREAlSaOTT

(/>
o'
3

(On# part p#r mlUlom)

CD

8
■
D

TABU u

TABIZ IB

SBOXS 145

SERH3 147A

(O '

i

RpophM#
3.

3
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CD

CD

■
D
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C
a
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3
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((//))

Hour#
0
1
S
8
4
8
6
7
6
9
10
11
IB
Tùtal
Treated

No.

No.

Iff ffiff
115
167
116
106
149
163
170
199
196
124
132
129

N#t#ph#—

40.4
69.7
46.6
42.2
64.8
64.4
63.2
67.9
96.2
48.6
48.4
47.2

1682 94.4

7?
68
46
69
77
47
99
64
44
67
71
82
72

i

Total

No.

?7t9 fl ? iil
24.5
18.1
27.8
30.7
20.4
20.9
25.8
18.8
23.8
28.0
30.0
27.2

760 24.6

fto p h # # #

P 0 # t4 le t# i.

Hour#

39.4
16.1
29.4
27.1
14.8
14.6
13.0
15.2
19.9
23.2
21.6
29.7

267
280
294
246
251
230
295
269
254
281
294
273
269

6
7
8
9
10
11
12

690 21.0

3092

Total
Treated

99
41
65
68
34
37
59
51
96
99
99
68

0
1
2
3
4

9

No.

117

109
129
104
122
190
119
175
132
150
128
130
138

k#t#phaa#

i

No.

90.9
60.8
47.3
90.0
95.8
99.2
66.9
97.4
95.1
54.9
38.6
62.7

7@
46
38
45
47
64
39
34
48
97
92
42
43

1964 96.4

P o 8 t4 i# t# $

No.

ye,3 1?

21.9
17.9
19.9
19.3
23.8
19.4
13.1
20.9
23.3
22.1
18.9
19.5

995 19.9

%

Tota]

27.6
21.2
33.2
30.7
20.4
21,4
20.4
21.7
23.7
23.4
22.5
17.7

214
212
220
244
269
201
260
230
245
235
222
220

695 23.6

2772

99
49
73
79
99
43
93
50
98
99
50
39

5

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TABIZ 1 (C0BtlnD»d)

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(On# p irt par m illion)

%
o
Z
J

1C

TABIZ IB

CD

8
■
D
C
5-

SRHS 143 AND 1474
Hour#

Prophma#
Bo. i

FSRdHIACBS COHVSRTED DSDiO BASE OF 50

O a tB D ttP

Total

134 26.6

509

0

43.7

50.0

29.6

50.0 26.6

50.0

32.0
18.5
29.1
28.9
17.8
17.6
16.6
17.5
21.7
23.3
22.0
22.1

494
466
468
495
499
454
529
464
526
489
495
485

1
2
3
4
5
6
7
8
9
10
11
12

44.9
63.5
47.0
46.1
59.9
62.1
64.8
62.7
54.8
51.5
52.9
54.2

51.2
72.4
53.6
52.6
68.3
70.8
73.9
71.4
62.5
58.7
60.3
61.8

23.1
18.0
23.9
25.1
22.2
20.3
18.5
19.8
23.6
25.2
25.1
23,7

39.0
30.4
40.4
42.4
37.5
34.3
31.3
33.5
39.9
42.6
42.4
40.1

60.2
34.8
54.7
54.3
33.5
33.1
31.2
32.9
40.8
43.8
41.4
41.5

3246 55.4 1313 22.4 1305 22.3

5864

0

m

45.7

149 29.6

CD

1

zsz

CD

Z

296
220
228
299
282
343
291
268
252
262
263

44.9
63.3
47.0
46.1
59.9
62.1
64.8
62.7
54.6
51.5
52.9
54.2

114
84
112
124
111
92
98
92
124
123
124
115

Hour#

3.

3
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C/)

3
4
5
4
7
8
9
10
11
IB
Total
Treated

Po#t4(ata.
Mataphaaa
Raal
N«u Raal Maw

Kataphmaa Foat-HaU.
Mo. % Mo. ^

23.1 158
18.0 86
23.9 136
25.1 143
22.2 89
20.3 80
18.5 88
19.8 81
23.6 114
25.2 114
25.1 109
23.7 107

Frophaaa
Raal Nam

32.0
18.5
29.1
28.9
17.8
17.6
16.6
17.5
21.7
23.3
22.0
22.1

TÂBLS 2

C
D
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PERCmmUZS

g

Q
.

FIOURSS
A. SHORT HOADONT

of total division

of

INDIVIDm STACKS

1$
—
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3
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(Fire parts per m illion)

"
D
C
D

3

TABU £A

C
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C
/
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o'

TABU £B

3

o
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C
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3
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C
D

3

%
o

3

Hours
0
1
£
3
4
5
6
7
8
9
10
11
1£
Total
Treated

aSRlSS 198
Frophaee Metashase PosWWta.
Total
No. *
lb ,
*
No. i
125 45.6 80 £9.9
£73
68 £4.9
£64
141 55.4 58 ££.0
65 £4.6
269
£09 77.£ £5 9.5 95 13.9
177 69.4
£55
57 2£.4 £1 8.2
120 53.8
£23
69 £8.3 40 17.9
129
29 22.5
60 46.5 40 31.0

In su ffieiea t DiTlsioo Figures

701 61.8 243 21.4

190 16.8

1134

Hours
0
1
£
3
4
5
6
7
8
9
10
11
12
Total
Treated

SERIES 192B
Pronhasa Nataphasa Fost4tota«
No. i
No. $
No. i
119 49.1 37 16.1 80 34.8
119 41.7
60 £2.1
98 96.2
166 71.2 30 12.7 98 16.1
38 14.8 33 12.9
185 78.3
47 £0.5
160 69.9
22 9.6
69 £1.4
80 £7.1
152 51.5
137 55.0 35 14.1
77 30.9
61 22.6
140 52.4 66 24.7

Total
£30
271
£36
256
229
295
249
267

In su ffieien t DlTislco Figures

1055 58.5 314 17.4 434 24.1

1803

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FSRCEKTACZS OF TOTAL DIVISION FIGGRSS OF DlDmDOAL STAGES

3

B. COmNDDUS TRSAH3ENT

CD

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TABU 6B

SKRUS 146

SmSS 151

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Hours
0
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2
3
4
5
6
7
8
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11. 12
Total
Trtattd

Prophatt
Mb.
117 48.1
146 53.1
166 63.6
142 61.7
117 62.9
126 56.3
119 57.8

Mtt&phwe PomWWW.
No. f
Mo.
Total
243
62 25.5 64 £6.3
275
73 26.5
56 20,4
47 18.0 46 18.4
261
43 18.7
230
45 19.6
52 27.9
17 9.1
186
72 32.1 26 U .6
224
67 52.5 20 9.7
206

la su fflo ltn t D ivision Figuras

816 59.0

354 25.6

212 15.3

1382

Prophm— Hfttphmmt Pomt^lmW»
Total
No.
No.
No.
269
106 39.4 82 30.5 81 30.1
242
69 28.5 60 24.8
113 46.7
304
159 52.3 70 23.0 75 24.7
226
54 23.9
135 59.7 37 16.4
274
161 58.8 66 24.1 47 17.1
302
55 18.2 44 14.6
206 67.2
209
13.9
123 58.8 57 27.3 29
206
135 65.5 42 20.4 29 14.1
158
105 66.5 23 14.5 30 19.0
Insuffieient Division Figuras

Hours
0
1
2
3
4
5
6
7
8
9, 10,
12
Total
%aatad 1134 59.0 419 21.8 368 19.1

1921

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Prophast

CD

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£68
£67
353
337
304
297
147
143
109

51.1
53.8
59.8
63.6
62.3
63.2
57.4
63.5
49.1

1957 59.7

20.4
28.2
20.2
20.4
22.9
23.2
30.1
24.9
34.2

149
89
118
85
72
64
32
26
37

28.5
17.9
20.0
16.0
14.7
13.6
12.5
11.5
16.7

524
496
590
530
488
470
256
225
222

797 24.3

423

15.9

3277

107
140
119
106
112
109
77
56
76

Hours
0
1
2
3
4
5
6
7
8

Fronhase
Real
51.1 50.0
53.8 52.7
59.8 58.6
63.6 62.3
62.3 61.0
63.2 61.9
57.4 56.2
63.5 62.2
49.1 48.1

Metanhase
Real Ne#
50.0
20.4
69.0
28.2
20.2 49.5
20.4 50.0
56.1
22.9
56.8
23.2
73.7
30.1
61.0
24.9
34.2 83.8

Poat-41eta.
Real Kern
50.0
28.5
17.9 31.5
20.0 35.0
16.0 28.0
14.7 25.7
13.6 23.8
12.5 21.8
11.5 20.1
16.7 29.2

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TABLE 9
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PERCmVKZS OF TOTAL DIVISION FIOORK OF IXDITIiJUAL STACSS
B. C0NTDID0D8 TSXAIUXNT
(Forty
8D
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SEBUS 95

CD

C

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TABU 9A

3
3.

p u rtt

Houri
0
1
z

3
4
5
6
7
8
9
10
11
U
Total
Tfoattd

Prophast liitftpbaM i^ost-Nota.
No.
No. f
No.
US 63.4 34 16.8 40 19.8
131 64.2 99 28.9 14 6.9
136 65.4 62 29.8 10 4.8
104 49.8 87 41.6
18 8.6
111 90.9 83 38.1 24 11.0
99 49.0 96 43.6 25 11.4
101 47.6 81 38.2 90 14.2
89 38.9 111 48.0 31 13.9
86 36.3 U6 95.2 29 10.9
89 24.2 122 99.7 44 20.1
86 36.1 U9 90.0 39 13.9
69 31.9 109 92.4 34 16.9
1061 44.1 1099 49.9

288 U.O

Total
202
204
208
209
218
220
2U
291
237
2U
238
208
2404

Hours
0
1
2
3
4
5
6
7
8
9
10
11
U
Total
»#ato6

Frophasa
NO.
137 91.7
U7 92.0
110 50.9
107 48.4
100 41.7
108 46.0
96 41.2
109 45.9
119 51.6
109 49.3
98 39,9
122 50.6
92 39.8

SERISS IBS
Natapbase Fost-Nata.
No.
Total
lA).
69 26.0
269
99 22.3
21 8.6
244
96 39.3
216
19 8.8
87 40.3
221
9 4.1
109 47.5
240
5.0
128 95.3
U
239
108 46.0 19 8.1
23
9.9
233
U4 48.9
231
90 39.0 36 15.6
223
77 34.9 31 13.9
221
2 0.9
110 49.8
248
139 93.6 17 6.9
241
9.0
107 44.4 U
231
129 59.8 10 4.3

1289 46.3 1284 46.1

211

7.6

2784

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table

FSRCSNTAG8S CONVERTED USU.G BASE OF 50

SSRUS 95 AKD 125 C013D3D
Hours
0
1
2
3
4
5
6
7
8
9
10
11
11
Total
Treatad

Aropkase
No,
^
265 56.7
858 57,6
246 58.0
811 49.1
211 46.0
807 45,5
197 44.3
194 48.0
801 43.7
168 36.8
184 37.9
187 41.6
98 39.8

xetapbaee Post-* eta.
No. >
No.
ft
95 19.9 109 23.3
155 34.6 35 7.8
149 35.1 89 6.8
198 44.6 87 6.3
211 46.0 36 7.9
204 44.8 44 9.7
55 11.9
195 49.8
801 43.5 67 14.5
56 18.8
809 44.1
238 58.7 46 10.4
258 51.8 50 10.3
46 10.8
816 48.1
10 4.3
119 55.8

8350 45.3 2339 45.1 499

9D

9.6

Total
467
448
424
490
456
456
445
468
460
440
486
449
831

hours
0
1
8
3
4
5
6
7
8
9
10
11
12

ProiAase
Real Lew
56.7 50.0
57.6 50.7
58.0 51.0
49.1 43.8
46.0 40.5
45.5 40.0
44.3 39.0
48.0 37.0
43.7 38.5
36.8 32.4
37.9 33.4
41.6 36.6
39.8 35.0

Metaphase
Real
New
19.9 50.0
34.6 86.8
35.1 88.1
44.6 111.9
46.0 115.5
44.8 112.4
43.8 109.9
43.5 109.2
44.1 110.7
52.7 132.3
51.8 130.0
48.1 120.7
55.8 140.0

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Real
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23.3
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7.8 16.8
6.8
14.6
13.5
6.3
17.0
7.9
9.7 20.9
11.9 25.6
14.5 31.2
12.8 26.8
10.4 22.4
10.3 22.1
10.8 21.9
9.8
4.3

5186

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Prophase
Hoars Count i
0
138 59.6
1
154 56.3
147 38.1
2
149 56.0
3
4
145 56 4
139 50.9
5
6
126 33.6
7
129 58 8
125 33.6
8
9
126 49.2
10
116 45.3
11
131 33.5
12
151 61.9
Total 1656 56.3

letapbase
Count
i
50 21.9
69 33.7
73 28.9
73 88.4
88 31.9
81 29.7
88 37.6
86 35.6
84 36.1
100 39.1
118 46.1
99 40.4
81 33.8
1056 35.0

F o st^ ta .
Count
^ Total
42 18.4 886
264
8.0
21
33 13.0 233
35 15.6 257
257
30 U .7
33 19.4 275
234
8.5
80
847
18.1
SO
84 10.3 833
856
30 U .7
856
82
8.6
6.1 845
15
4.9 844
18
385 10.9 3019

Hours
0
1
2
3
4
5
6
7
8
9
10
U
12

Propbase
Real Nee
59.6 50.0
56.3 48.9
58.1 48.7
56.0 48.7
56.4 47.3
50.9 48.7
58.6 45.1
58.2 43.8
53.6 45.0
49.2 41.3
45.3 38.0
53.5 44 9
61.9 51.9

dbins bass of so

Metaphase
Real New
21.9 50.0
33.7 76.8
28.9 65.9
28.4 64.8
31.9 72.7
29.7 67.7
37.6 85.7
35.6 81.2
36.1 82.3
39.1 89.1
46.1 105.1
40.4 92.1
33.8 75.7

Post-Meta.
Real New
18.4 50.0
8.0 21.8
13.0 35.4
13.6 37.0
U .7 31.8
19.4 52.8
8.5 23.1
12.1 32.9
10.3 28.0
U .7 31.8
8.6 23.4
6*1 16.6
4.9 13.3

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PKRCBTPAOagS OF TOTAL DIVISION FIGURES OF INDIVIDUAL STACKS
D. UNSEATED

CD

(/)
C/)

(Root tip# oolleoted at oonateutlra hour#)
CD

8
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Hour#
0
1
2
5
4
5
6
7
8
9
10
11
12
Total

TABZK ISA

TABIK 13B

SKRUS 127

SERIES 132

Fropha## Nataphas# Poat-ICaU.
No. i
No. i
Bo. f
97 26.7
106 90.5 49 22.9
113 48.1 49 20.8
75 31.1
93.7
48
96
25.5
21,0
123
141 59.1 49 17.6 70 27.5
102 44.1
94 25.4
79 32.5
98 41.3 44 18.6
99 40.1
123 47.1 68 25.3
72 27.6
UO 49.8 62 26.0 49 22.2
117 52.5 82. 25.3
54 24.2
97 23.8
119 47.9 68 26.3
107 46.7
91 22.3 71 51.0
101 44 5 97 25.1 69 30.4
116 48.5 93 21.8 72 29.6
1476 48.5 708 23.2 862 28.3

Total
214
255
229
296
251
237
261
221
229
240
229
227
243
3046

Hours
0
1
2
3
4
5
6
7
8
9
10
11
12
Total

Frophaaa Mataphaaa PostData.
No.
f
No. a
No.
i
122 48.6
64 25.5 65 25.9
112 41.2 70 25.8 90 35.1
109 49.4 61 25.4 70 29.2
120 52.1 96 25.2 52 22,7
62 24.0
112 43.5 84 32.6
98 46.2 99 27.8 55 25.9
125 52.1 63 26.3 52 21.7
65 26.1
55 22.8
123 51.0
116 38.9 85 28.5 97 32.5
59 25.4 67 28.9
106 45.7
91 39.6
69 30.0 70 30.4
51 32.5 35 22.3
71 45.2
1505 45.6 786 27.5 770 26.9

Total
251
272
240
230
298
212
240
241
298
232
230
157
2861

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TABIZ 14

PSÏCKNTAC3S OF TOTAL DIVISION nClHES OF INDIVIDUAL STAGES
D.UNTREAISD

■
O
C
D
%

(Root tip s from same onion at sams hour)

o
=
5

TABIZ I t t

TABIZ 14A

8
T3

FERCENTAGSSHCB
Oaxm
NVERB
Ta^sassB
ED USING
BASS OF 50
SBBxasssa^Bse

SERIES 137B

C
5-

Prophase

Metaphase

C
D

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c
3.

3"

C
D

cB
3

■
D
Q
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C
a

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3
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D
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3

No.

2
236
3
238
4
211
231
6
7
237
229
e
232
10
11
254
IS
195
13
220
15
220
21
213
TPtal 2716

Count

Count

Count
54.4
47.8
51.1
54.7
52.9
50.9
54.1
59.2
44.1
47.9
49.9
43.4
50.7

94
128
124
99
111
105
109
81
111
125
105
165
1355

21.7
25.7
30.0
25.5
24.8
23.3
25.4
18.9
25.1
27.3
23.8
33.2
£5.3

Prophase

PostData,

104
132
78
92
100
116
88
94
136
114
U6
115
1285

ft Total
24.0 434
26.5 498
18.9 413
£1.8 422
£2.3 448
25.8 450
20.5 429
21.9 429
30.8 442
£4.8 459
26.3 441
23.4 491
23.5 5556

Root
No.

Count

2
3
4
6
7
8
10
11
12
13
15
21

54.4
47.8
51.1
54.7
52.9
50.0
54.1
59.2
44.1
47.9
49.9
43.4

Metaphase

Count

Count
50.0
44.0
47.0
50.5
48.7
46.0
49.8
54.5
41.3
44.1
45.9
39.9

21.7
25.7
30.0
23.5
£4.8
23.3
25.4
18.9
25.1
27.3
23.8
33.2

Poat-Msta.

50.0
59.1
69.0
54.0
57.0
53.6
58.4
43.5
57.7
62.8
54.7
76.4

24.0
26.5
18.9
21.8
22.3
25.8
20.5
21.9
30.8
24.8
26.3
23.4

i

50.0
55.1
39.3
62.0
47.4
53.7
42.6
45.6
64.1
51.6
54,7
48.7

S

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140
TA B U 16
AVKRAGB NCM
BKRS OF DI7ISI0N FIGURES PER AXIS'
A. SHORT TBBAHŒNT
(On# p#rt per m illion)

Hoars
0
1
£
3
4
9
6
?
8
9
10
11
IS
Total
Treated

ssRxsS 143
Count
Axes

ÜBKIXS 147A
Count
Axes

COMBINED
Count
Axes

Arg.

£67
£80
£84
£48
£51
£30
£55
£69
£54
£81
£54
£73
£65

15
14
7
3£
1£
31
£0
1£
9
7
6
9
8

£36
£14
£1£
£80
£44
£69
£01
£60
£30
£45
£55
£££
££0

£7
£8
17
IT
19
18
19
£0
8
16
18
£3
£6

SCO
494
466
468
498
499
454
589
464
586
489
495
485

40
48
£4
49
31
46
59
38
17
83
£4
38
34

18.6
11.8
19.4
9.6
16.0
10.9
11.6
16.5
£7.3
88.9
£0.4
15.9
14.5

309£

167

£77£

£86

9864

393

14.9

TAELS IT
A.VKHAÆ
K Nin

OF D IV ISIO N FIGURES iV R AXIS
A, SHORT TRSAWSNT

(Five parts per m illion)

Hours
0
1
£
3
4
5
6
7
Total
Treated
1.

SKRUS 136
Axes
Count
IT
£73
14
£64
IS
£63
£4
£55
36
££3
41
1£9

1134

1£7

SERIES 15£B
Count
Axe#
£30
11
10
£70
£0
£56
£56
14
££9
33
£95
19
£8
£49
£9
£67
1803

147

CQUBDDD)
Count
Axe#
£8
503
£4
555
38
499
38
511
69
458
60
484
£8
£49
£9
£67
£937

£74

18,0
£8.3
19.6
13.5
6.6
7.1
11.3
9.8
10.7

The term "mzle" em need in Tables 16 through £80 re fer to the eoaseeutir e series of high powered fie ld s taken from edge to edge of the
corersllp*

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

141
T4BLS 10
AVXRAOB NIKBBRS OP DIVISION PIODRXS PER Ayra
A. SHORT TRIATNENT
p arts par m illion)

Hours
0
1
2
3
4
5
6
Total
TYsatsd

SERIES 92
Count
Axas
207
12
229
9
201
12
209
12
199
31
209
19
127
40
1164

129

SERIES 140
Count
Axes
261
10
257
10
290
16
245
20
228
35
212
31
—

1232

-

112

OORBINED
Oount
Axes
Atc*
468
22
21.3
482
19
25.4
491
28
17.5
448
32
14.0
427
66
6.5
421
50
8.4
127
40
3.2
2396

235

10.2

TABU ;19
AVERAOBK NTKBERS 07 DIVISION FIGURES PER AXIS
A. SHORT TREATMENT
(Forty parts per million)

Hours
0
1

2
3
4
5
Total
Trsated

sraiBs 96
Count
Axes
17
209
17
214
13
215
230
14
28
201
17
69
929

89

SERIES 120
Count
Axes
IB
24?
209
29
206
42
24
241
161
44
28
18
845

157

OCKBINBD
Count
Axes
Ats.
15.0
35
456
9.2
423
46
7.7
55
4SI
12.4
471
38
5.0
72
362
2.8
97
35
1774

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

246

7.2

14S
TAHLX SO
ATHRAGZ NUkBSRS OF DIVISION FIGURES FKR AXIS
A. SHCRT IREATKXNT
(Eighty parts par m illion)

Hours
0
1
2
3
4
5
6
Total
Treated

SERIES 118
Count
Axes

SmiES 122
Count
Axes

£51
209
268
260
218
211
216

21
14
13
17
25
29
43

297
221
310
256
288
163

1382

141

1178

-

17
19
13
14
19
24
-

89

COMBINED
Count
Axes
Avg.
548
430
578
516
446
374
216

38
33
26
31
44
53
43

14.4
13.0
28.2
16.6
10.1
7.1
5.0

£560

230

11.1

TABLE 21
AVERAGE HUkBSRS OF DIVISION FIGURES PER AXIS
B. CONTINUOUS IREATKENT
(One part per m illion)

Hours
0
1
2
3
4
5
6
7
8
Total
Treated

SERIES 146
Oount
Axe#
245
275
261
230
186
224
206

OGHBINB)
Axe#
Oount

AVg.

20
15
16
25
33
31
41

269
248
304
226
274
302
209
206
158

8
18
8
10
14
19
30
26
35

512
517
565
456
460
526
415
206
158

88
33
26
35
47
50
71
26
35

18.3
15.7
21.7
13.0
9.8
10.5
5.9
7.9
4.1

163

1921

160

3303

323

10.2

«
«

1139

SERIES 151
Count
Axe#

Reproduced with permission o f the copyright owner. Further reproduction prohibited without permission.

TABLE 22
AVERAGE NUMBERS 07 DIVISION FIGURES PER AXIS
B. OONTIiaJOXJS IREAI&3NT
(Flv# parts par million)

Hours
0
1
2
3
4
5
6
7
9
Total
Traatad

SERIES 139
Azas
Count
267
18
286
19
279
11
290
19
215
28
213
SO
—

—
—

1199

107

SERIES 182A
Count
Axas
297
6
240
10
311
14
300
14
275
20
297
17
256
16
27
225
S»
96
2084

194

OOKBIKED
Count
Azas
524
24
496
29
590
25
930
35
48
488
470
47
256
16
27
225
36
222
5277

261

Ate.
21.8
17.1
23.6
16.1
10.2
10.0
16.0
8.3
6.2
12.6

TABLE 29
AVERAGE NULIBERS OF DIVISION FIGURES PER AXIS
B. CONTINUOUS TREATMENT
(Tmanty parts psr m illion)

Hours
0
1
2
3
4
5
6
7
8
9
10
11
12
Total
Traatad

SERIES 128
Count
Azas
230
16
232
10
232
8
228
IS
236
13
17
221
16
249
20
236
18
239
245
20
12
216
16
222
16
219
2769

179

SERIES 194
Oount
Azas
8
213
220
6
268
8
276
6
10
256
277
9
272
11
275
9
295
11
249
8
256
10
248
8
242
11
3088

107

COkBINED
Atz.
Count
Azas
18.9
443
24
28.3
452
16
31.3
500
16
26.5
19
904
21.4
29
492
19.2
26
498
19.1
27
515
17.6
29
511
17.0
29
492
17.9
28
490
22
21.5
472
19.6
24
470
17.1
27
461
9857

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

286

20.9

TABIÆ £4
AVZ&AC3 UXA1BER3 07 DIVISION 7I0URBS PSR AXIS
B. CONTINUODS TREAHCENT
(Torty part# par m illion)

Hour#
0
1
2
3
4
5
6
7
8
9
10
11
12
Total
"Rraated

SXRISS 95
Count
Axe#
SOS
IB
204
17
806
11
209
9
216
17
220
15
212
14
22
231
237
16
219
23
238
21
208
30
-

2404

-

195

SERIES 125
Count
Axa#
19
265
23
216
13
221
17
240
17
235
16
21
233
231
19
223
16
221
23
248
19
241
26
231
18
2784

228

COMBINED
Count
Axa#
467
37
448
40
424
24
430
26
458
34
455
31
445
55
462
41
460
32
440
46
40
486
449
56
231
18
5186

423

At#.
12.6
Ü .2
17.7
16.5
13.5
14.7
12.7
11.3
14.4
9.6
12.2
8.0
12.8
12.3

TABLE 25
AVSRAGX NUkBRS 07 DIVISION 71GORES PER AXIS
B. COHTTNUOOS TREATMENT
(Eighty part# par m illion)

Hour#
0
1
2
3
4
S
6
7
8
Total
Treated

SERIES 102
Count
Axes
206
14
212
23
20
214
231
12
234
26
29
214
_

SERIES 115
Count
Axa#
18
221
245
17
221
15
15
281
22
269
-

—

212
163

39
33

261
237
260

11
13
16

1480

182

1774

109

COMBINED
Count
Axa#
32
427
40
457
35
435
27
512
503
48
29
214
11
261
52
449
49
423
3254

Reproduced with permission o f the copyright owner. Further reproduction prohibited without permission.

291

Arg.
13.3
11.4
12.4
19.0
10.5
7.4
23,7
8.6
8.6
11.2

140

lABUE 86
AVSRAGEB NUMBERS OF DIVISION FIGURES ITO AXIS
0. UNTREATED
(Root tip a oolloetad a t eoesaautlre hours)

Hours
0
1
2
3
4
5
6
7
8
9
10
11
12
Total

SERIES 127
Count
Azss
214
13
238
18
229
13
256
13
231
12
237
13
261
14
221
12
223
18
240
19
229
16
227
14
243
19
3046

SERIES 158
Count
20
251
272
13
240
9
230
11
288
15
212
5
240
11
241
13
298
8
232
10
230
18
187

32

465
807
469
486
489
449
801
462
821
472
489
227
400

2861

168

8907

-

191

OOfSINED

-

TABIE 27
AVTOAGE NTEIBERS OF DIVISION FIGURES PER AXIS
C. XDnREATED
(Root tip s oollsotsd from sans onion at same hour)
Hours
0
1
2
3
4
5
6
7
8
9
10
11
12
Total

Root No.
2
3
4
6
7
8
10
11
12
13
15
21
—

Oount
434
498
415
422
448
480
429
429
442
459
441
491
-

5356

Axas
21
21
25
25
24
12
24
22
14
19
33
24
-

262

AVg.
20.7
23.7
16.8
18.4
18.7
37.8
17.9
19.5
31.6
24.2
13.4
20.5
—
20.4

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

33
28
22
24
27
18
25
25
26
29
34
14
51

l4.1
18.1
21.3
20.3
18.1
24.9
20.0
18.8
20.0
16.3
13.5
16.2
7.8

386

16.6

IM
TABLE 8 8
AVERACHK NIHBKRS OF DIFISZON FIGURES PER AXIS^
TABIE 88A
ZERO HOURS OF TREATED BULBS
Part# par n illlm i

Count

Axas

Arg*

1
S
80
40
80

903
903
468
496
948

40
88
88
39
38

18.6
18.0
81.3
19.0
14#4

Total

8478

163

15.8

TABIE 88B
3CR0 HOURS OF TREATED BULBS
CONTINUOUS TREATMENT
Part» par m illion

Count

Axas

Avg.

1
8
80
40
80

518
584
443
467
487

88
84
84
37
38

18.3
81.8
18.9
18.6
13.3

Total

8373

149

16.4

TABIE 880
SRHMAXT OF ALL UNTREATED MATERIAL

0-Bonr Short
0-Boor Continuous
ConaaeutiTo Hours
Sam# Hour
Total
X*

Count

Axas

Arg.

8478
8973
9907
9396

163
149
396
868

19.8
16.4
16.6
80.4

16114

986

17.4

Th# %«m "mzlm" a# uaad in Tabla» 16 throagb 880 rafar to tha
eoaaaeatlTa aarla# of high poaarad fiald a takan from #dga to
adga of tha oorarallp*

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

1 4 !f
TàBLB 2 9
HOURLY CHA^'^ES IN AVZRACg NI&.BER OF DIVISION FIGURES PER AXIS
SHORT raSATll®iT

Hr#.
0
1
S
3
4
5
6
7
8
9
10
11
12

1 P. P.M.*
Arg. Dev.

9 P.P.M.
Avg.

18.0
0.0
12.6
22.3
11.8
-0.8
6.8
15.6
19.4
13.5
9.6 - 3.0
6.6
16.0 + 3.4
7.1
1C.9 - 1.7
11.3
11.6 - 1,0
9.2
16.5 ♦ 3.9
+14.7
27.3
22.9 +10,3
20.4 + 7.8
15.5 + 2.9
14.3 + 1.7
* Part# per m illion

Dev.
0.0
+ 4.3
- 2.4
—4.5
-11.4
-10.9
- 6.7
- 8.8

20 P.P.M.
Avg. Dev,
21.3
25.4
17.5
14.0
6.5
8.4
3.2

0.0
+ 4.1
- 3.8
—7.3
-14.8
-12.9
-18.1

40 P.P.M.
Avg.

Dev.

13.0
9.2
7.7
12.4
5.0
2.8

0.0
- 3.8
- 5.3
—0.6
- 8.0
-10.2

80 P.P.M.
Avg. Dev.
14.4
13.0
22.2
16.6
10.1
7.1
9.0

♦
♦
-

0.0
1.0
7.8
2.2
4.3
7.3
9.4

TABLE 30
HOURLY CHANGES IN AVERAGE NIABER OP DIVISION FIGURES HER AXIS
CONTINUOUS TREATMENT

Hr#.
0
1
£
9
4
9
6
7
8
9
10
11
12

1 P. P.k.*
Arg. Dev.
18.3
15.7
21.7
13.0
9.8
10.9
5.9
7.9
4.1

0.0
- 2.6
+ 3.4
—9.3
—8.9
- 7.8
-12.4
-10.4
—
14.2

5 P. P.M.
Avg. Dev.
21.8
17.1
23.6
16.1
10.2
10.0
16.0
8.3
6.2

0.0
- 4.7
+ 1.8
- 9.7
-11.6
-11.8
- 5.8
—
13.5
—
15.6

80 P.P.k.
Avg. Dev.

40 I».P.M.
Avg.
Dev.

80 P.P.M.
Avg. Dev.

18.9
28.9
31.3
26.9
21.4
19.2
19.1
17.6
17.0
17.9
81.9
19.6

12.6
11.2
17.7
16.9
13.9
14.7
12.7
11.3
14.4
9.6
12.2
8.0
12.8

13.3
11.4
18.4
19.0
10.9
7.4
83.7
8.6
8.6

0.0
+ 9.8
+18.8
+ 8.0
+ 2.9
+ 0.7
+ 0.6
- 0.9
—1.5
—1.0
+ 3.0
+ 1.1
- 1.4

0.0
- 1.4
+ 9.1
+ 3.9
+ 0.9
+ 2.1
+ 0.1
- 1.3
+ 1.8
- 3.0
—0.4
—4.6
+ 0.2

Part# par m illion

Reproduced with permission o f the copyright owner. Further reproduction prohibited without permission.

0.0
- 1.9
- 0.9
+ 9.7
- 2.8
- 9.9
+10.4
- 4.7
- 4.7

UM

TABU 51
HOtJRLY CKAKOBS IN AVBRACg NUBBR OT DIVISION FIGURES PER AXIS
TJNIRSAnED

(Root tip # tmk#n a t oonsootitlr# hour#)

Hour#
0
I
2
5
4
6
6
7
e
9
10
11
12

Arg#

Dot.

14.8
18.9
21.5
20.5
18.1
24.9
20.0
18.5
20.0
16.5
15.5
16.2
7.8

0.0
♦ 4.1
* 6.5
+ 5.5
+ 3.5
♦10.1
♦ 5.2
♦ 5.7
♦ 5.8
♦ 1.5
—1.5
♦ 1.4
- 7.0

TABU 32
HOURIY^ CHANGES IN AVXRACB NOiHCR OF DIVISION FIGURES VSR AXIS
UNTREATED
(Root tip# takan from sam# onion a t aam# hour)

Root No#
2
5
4
8
7
8
10
11
12
15
15
21

Avg.

Dot.

80.7
83.7
16.5
ie .4
18.7
37.5
17.9
19.5
51.6
84.2
13.4
20.5

0.0
♦ 3.0
—4.8
—2.3
—8.0
♦16.8
- 2.8
—1.8
♦10.9
♦ 3.5
- 7.5
—0.8

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

149

TABIX 3S
IVFCSiTACOS OF AEBRRAM
T HtOHlASSS
A. SHORT TREATMENT
(On# part par million)

SERIES 143
Abar* Total
%
108 10*7
11
19
113 16.8
19
167
7.8
9
116
4.3
14
106
3.8
6
149
4.0
19
165
9.8
3
170
1.8
8
199
1.9
198
4
2.9
9
184
4.0
1
132
0.8
8
129
1.6

Hrs*
0
1
s
3
4
9
6
7
8
9
10
11
12
Total
Treated

89

1682

9.3

SERIES 147A
Abar. Tbtal
0
117
3
109
8
189
1
104
0
188
0
190
0
119
1
173
0
138
1
130
188
4
1
130
1
138
14

1964

36
0.0
8.8
1.6
1.0
0.0
0.0
0.0
0.6
0.0
0.8
3.1
0.8
0.7
0.9

OOMBINED
Abar. Total
11
880
28
888
19
896
880
6
14
888
6
899
19
888
4
343
8
891
9
288
9
888
868
2
863
3

36
9.0
9.9
0.1
8.7
6.1
8.0
9.3
1.8
0.7
1.7
8.6
0.8
1.1

3246

3.2

OOUBIHB)
Abar. Total

36

103

TABIE 34
nCRCSRTACaCS OF ABERRANT IROFHASBS
A. SHORT TREATMENT
(FIt# parts par million)

SERIES 138
Hr8.

Abar.

0
0
1
8
2
73
3
116
4
109
9
96
6
7
Total
Treated 390

Total

SERIES 19BB

%

Abar.

Total

%

189
141
208
177
180
60

0.0
1.4
36.0
69.5
89.8
93.3

6
4
7
94
106
74
79
72

113
113
168
189
160
198
137
140

9.3
3.8
4.8
50.8
66.3
48.7
57.7
51.4

6
6
80
210
809
130
79
72

701

49.9

436

1059

41.3

786

Reproduced with permission o f the copyright owner. Further reproduction prohibited without permission.

8.5
838
8.4
894
371 81.6
368 58.0
880 74.6
218 61.3
137 57.7
140 51.4
1756

44.8

150

TABLE 3 5
PERCBITTAOrnS 0 7 ABBOURT HtOFHASXS
A. SHORT TREAMNT
(Twenty pnrts per million)

Hra*

SERIES 98
Abar# Total

0
7
1
14
£
38
3
84
4
47
5
71
6
38
Total
Treated 286

SBtlES 140
Abar. Total

St

113
148
186
114
138
146
94

6.8
9.9
85.4
81.1
35.6
48.6
40.4

0
9
84
174
158
90

754

30.0

515

188
145
164
185
163
98

-

-

753

%

COMBINED
Abar. Total

0.0
6,3
51.8
94.1
96.9
91.8
-

68.4

%

7
85
116
198
805
161
38

841
885
890
899
895
844
94

8.9
8.1
40.0
66.8
69.5
66.0
40^4

741

1507

49.8

TABUS 36
FEPCENTACBES OF ABERRANT FROHUSBS
A. SHORT TREAMNT
(Forty porta per m illion)

Hrs.

SERIES 96
Abar. Total

0
15
1
13
8
41
3
61
4
75
5
86
Total
Traatad 816

%

SERIES 180
Abar. Total

St

OOKBIHED
Abar. Total

St

110
185
105
114
149
48

13.6
10.4
39.0
53.5
50.3
54.8

84
77
90
117
101
81

186 19.0
118 68.8
109 88.6
130 90.0
108 93.5
81 100.0

39
90
131
178
176
47

836
837
814
844
257
69

16.5
38.0
61.8
73.0
68.5
68.1

541

59.9

406

480

688

1081

60.9

84.6

Reproduced with permission o f the copyright owner. Further reproduction prohibited without permission.

101
TABLS S7
FSROSNTAÛBS OF ABDtRAKT IftOFHASBS
A. SHORT TREAH3NT
(Eighty parts psr m illion)

Hrs.
0
1
2
3
4
5
6
Total
Traatad

SERIES lie
Abar. Total

SBtTMS 188
%

Abar. Total

S9
64
80
134
118
164

13?
6,1
187 28.8
150 48.7
124 66.5
136 98.5
124 95.2
164 100.0

31
84
125
118
187
117

094

885

565

-J ft

154
146
168
186
139
127

—

72.0

%
20.1
57.5
77.8
88.9
91.4
98.1

-

700

-

80.7

OOMBINED
Abar. Total
39
113
189
197
861
235
164
1159

%

886 13.6
273 41.4
318 60.6
850 78.8
875 94.9
851 93.6
164 100.0
1585

76.0

TABLE 38
PERCEKTAOES OF ABRRANT HtOFHASSS
B. CONTIKÜCÜS
(Ona part par m illion)

Hrs.
0

1
8
3
4
5
6
7
8
Total
TYaatad

Abar, Total
4
36
27
80
92
86
98

117
146
166
148
117
126
119

1183

Abar.

Abar. Total

Total

—

1
24
38
138
88
117
97

106
113
159
135
161
203
183
130
105

1.9
0.0
0.6
17.0
19.9
60.0
71.5
86.7
98.4

6
36
88
109
124
218
186
117
97

883
259
385
877
278
389
248
135
105

8^7
15.9
8.6
39.4
44.6
66.3
76.9
86.7
98.4

37.8

491

827

59.4

915

1950

46.9

3.4
84.7
16.3
59.9
78.6
68.3
88.4
—

424

COMBINED

SERIES 151

SERIES 146

8

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

m

TABUS 39
FEHCSKTAGaiS 07 ABIRRAKT HIOFHASBS
B. CONTINÜODS HŒA*MnJT
(five parts psr m illion)

Hrs.
0
1
2
3
4
5
6
7
8
Total
% sats4

SERIES 139
Abar. Total
5
141
29
146
121
168
138
154
119
129
134
136
-

-

941

3.5
19.9
76.6
89.6
92.2
98.9
—

783

74.8

SERIES 192A
Absr. Total
A
9
127
7.1
121
0.8
1
96
199 49.2
169
183 92.3
171
175 97.7
161 92.9
149
137
147 93.2
143 99.3
142
107
109 98.2
972

1234

COMBINED
Absr. Total
14
268
30
267
217
353
307
337
290
304
283
297
137
147
142
143
107
109

5.2
11.2
61.5
91.1
95.4
99.3
93.2
99.3
98.2

1513

1957

77.3

2.7
53.8
68.0
78.5
71.7
79.0
88.0
97.3
93.2
91.3
92.8
97.9
92.9

COIdBINED
Absr. Total
217
35
15t
264
295
226
217
275
266
202
278
225
254
222
249
234
234
224
229
215
240
223
243
236
229
218

16.1
&7.6
76.6
78.9
75.9
80.9
87.4
94.0
99.7
93.9
92.9
97.1
95.1

83.7

2594

3056

84.f

78.8

TABLE 40
PSRCEKTAGES OF ABERRANT ÎROFHASES
B. CONTINUOUS TREATNENT
(Twsntj parts par million)

Hrs.
0
1

2
3
4

5
6
7

a
9
10

n
12
to ta l
Trsatsd

SBIXXS 128
Absr. Total
32
105 30.9
132 61.4
81
142 85.9
122
112 79.5
89
114 81.6
93
111 83.8
93
88 86.4
78
106 89.3
92
87 100.0
87
103 97.1
100
87 93.1
81
96.1
105
99
88 96.9
87
1100

1270

86.6

SERIES 134
Absr. Total
112
3
7i
132
193
104
128
163
109
192
167
132
166
146
142
146
147
137
115
126
142
193
137
140
131
141
1494

1786

%

Reproduced with permission o f the copyright o w n e r Further reproduction prohibited without permission^

Jt

105
TABIX 41
HKCKNTAGHES OF ASBRRAKT HiOFHASSS
B. OONTIKUOOS
(Forty porta par m illion)

Hra.
0
1
2
3
4
9
6
7
8
9
10
11
12
Total
Traatad

SBRIX8 99
Abar. Total 7
128
9.9
26
191 19.8
90
196 22.1
49
104 47.1
44
111 99.6
99
99 93.9
46
101 49.5
77
89 86.9
83
86 96.9
91
99 96.2
86
86 100.0
69 98.9
64
609

97.4

1061

SERIES 129
Abar. Total
Â
19
137
9.9
127 19.7
29
110 80.9
89
99
107 92.9
100 99.0
99
106 98.1
106
96 99.0
99
104
109 99.0
119
119 100.0
104
109 99.4
99
98 94.9
122
122 100.0
92
92 100.0

OOKBINED
Abar. Total
20
269
7.9
91
298 19.8
119
246 48.4
148
211 70.1
143
211 67.8
199
207 76.8
141
197 71.6
181
194 93.3
198
201 98.9
199
162 99.7
179
184 97.3
186
187 99.5
92
92 100.0

1143

1792

1289

88.7

2990

74.6

OOKBIBED
Abar. Total
616
264
7Î
272
107
277
169
264
184
110
92
98
89
188
196
160
149

17.6
28.4
39.3
61.0
69.7
47.3
86.7
83.0
90.6

1633

59.6

TABIE 42
FEHCENTA(2S OF ABERRANT HtOFHASSS
B. OONTIKÜO^ TREAIkENT
(Eighty parta pmr m illion)

Hra.
0
1
2
3
4
5
6
7
8
Total
Ifeated

SERIES 102
Abar. Total
7
93
127
l i
156
40
117
99
127
70
110
92
«

67
59

99
66

362

798

7.9
11.8
29.6
50.4
55.1
47.3
—
70,5
89.4
45.4

SERIES 119
Abar. Total
96
119
137
éà
67
116
110
160
137
114

65.6
43.8
57.8
68.8
83.2

-

-

—

89
89
86

98
93
94

86.7
99.7
91.9

611

835

73.2

973

Reproduced with permission o f the copyright owner. Further reproduction prohibited without permission.

154

TABIS 45
PgRCKNTACgS OF ABERRANT HtOFHASES
C. IinXRIdlTTANT TRIAHCSNT
(Forty parts per m illion for 15 minutes every teo hours)

SERIES 186
Hours
0
1
8
3
4
8
6
7
8

Abar.
6
41
56
109
144
143
165

Total

Total
96
117
113
110
147
145
166

4
6,1
35.0
49,6
93,6
98.0
98,6
99.4

65

65

100,0

717

863

93,7

-

TABU 44
PERCENTAŒS OF ABERRANT FHOFHASES
C• INTER&ITTANT TREAB/KNT
(Forty parts per m illion for IS minutas every hour)

Hours
0
1
8
5
4
5
6
7
8
9
10
11
18
Total

Abar,

SERIES 135
Total

1
47
186
138
131
186
181
187
185
184
113
189
151

136
154
147
149
145
139
186
189
185
186
116
131
151

4
0,7
30.5
85.7
88.6
90.3
90,6
96.0
96,4
100.0
98,4
97.4
98,5
100,0

1458

188#

96,7

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

180
45
T^ROWTAŒgS OF ABIBRANT mOW ASÈS
D. UNTREATED
(Root tips oollsotsd st oonssoutiTs hours)

Tîra.
0
1
2
3
4
5
6
7
8
9
10
11
12
Total

sroiBS 127
Absr. Total
8
106
1
113
4
123
10
141
8
102
6
98
123
4
4
lie
5
117
6
115
107
5
6
101
118
3
60

1476

SERIES 158
Absr. Total
À
14
122 11.5
11
112
9.8
5
109
4.6
10
120
8.3
6
112
5.4
9
98
9.8
11
125
8.8
7
123
5.7
4
116
3.4
7
106
6.6
9
91
9.9

2.8
0.9
3.6
7.1
7.8
6.1
3.3
3.6
4.3
5.2
4.7
5.9
2.8

-

4.4

-

-

7

71

9.9

100

1183

8.8

G0> BillED
Absr. Total
17
230
12
225
9
232
20
261
14
214
196
15
15
248
11
233
9
233
221
13
14
198
6
101
10
189
165

8781

4
7.4
5.3
3.9
7.7
6.5
7.7
6.0
4.7
3.9
5.9
7.1
5.9
5.3
5.9

TABU 46
PERCENTAGES OF ABERRANT IROFHASES
D. UmHBAUO
(Root tips oollsotsd froBi ssms 'OhIon a t sans hour)

1
Hrs.
0
1
2
3
4
5
6
7
8
9
10
11
18
Total

Root No.
8
3
4
6
7
8
10
11
12
13
15
21
-

Absr.

Total

6
10
1
2
5
9
6
8
8
2
3
0

836
838
211
831
837
889
838
854
195
220
220
213

8.8
4.2
0.5
0.9
8.1
3.9
8.6
0.8
1.0
0.9
1.4
0.0
-

8716

1.8

48

Reproduced with permission o f the copyright owner. Further reproduction prohibited without permission.

106
lABUE 47
HCRCSNTACSS Of ABERRANT HIOPHASKS
TABLE 47A
aRO H00R8 OF TREATED BULBS

P srta par m illion

Abar*

Total

ft

1
5
SO
40
80

11
6
7
39
39

SSO
838
841
836
886

5.0
8.5
8.9
16.5
13.6

Total

lOS

ISSl

10.8

TABLE 47B
2ZR0 HOURS OF TREATED BULBS
CONTINUCUS TREATMENT
P arts par m illion
1
5
SO
40
80

Abar.

Total

6
14
35
SO
37

883
868
817
865
812

8.7
3.8
16.1
7.5
17.5

1185

9.5

US

Total

ft

TABLE 470
SWkAKY OF ALL UNTREATED MATERIAL
Abar.

Total

ft

0-Hour Short
0«Hour Contlnwma
ConaaoutlTa Boura
Sama Hour

lOS
US
165
48

1881
1185
8781
8716

10.8
9.5
5.9
1.8

Total

4S7

7903

6.6

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

19T
TABU 46
FEBCSNTACaSS OF SCATTSRS) UXTAPHA3ES

A . SHORT ntXAlUXNT
(On# p#rt p#r m illion)

arm.
0
1
2
3
4
5
6
7
8
9
10
11
12
Total
TYoated

SBRIXS 145
Soatt* Total
0
75
1
68
0
46
1
69
5
77
1
47
0
55
s
64
0
44
4
67
71
8
0
82
72
4
21

760

~

0.0
1.8
0.0
1.4
5.9
2.1
0.0
5.1
0.0
6.0
7.0
0.0
5,5
2.8

SERIES 147A
Soatt. Total
a
0
76
0.0
1
2.2
46
2
38
5.3
0
45
0.0
0
47
0.0
0
0.0
64
0
39
0.0
2
34
5.9
2.1
1
48
0
57
0,0
52
1
1.9
42
1
2.4
7.0
43
3
11

553

1.8

O
C
X
ibxnbd
Soatt. Total
0
149
2
114
2
84
1
112
3
124
1
111
0
92
4
98
92
1
124
4
6
123
1
124
7
115
32

'

________

0.0
1.8
2.4
0.9
2.4
0.9
0.0
4.1
1.1
5,2
4.9
0.8
6,1

1313

2,2

GOKBIEED
Soatt. Total
117
0
116
4
55
8
21
95
85
41
27
103
35
1
4
66

0.0
3,4
14.6
22.1
48,2
26.2
2.9
6.1

557

19,0

•

TABIE 49
FERCENTACaCS OP SCATTERED liETAFHASES
A. aaORT TREATMWT
(FIt# parta par million)

Hr#,
0
1

2
3
4
5
6
7
Total
Traatad

SERIES 156
Soatt* Total
80
0
4
7
21
35
22

58
25
57
65
40

li
0.0
6.9
28.0
36.8
52.4
55.0

—
«

87

•

243

—

35.8

SERIES 152B
Soatt. Total
0.0
37
0
60
0.0
0
3.3
30
1
0.0
38
0
56.4
22
6
7,9
63
5
2.9
35
1
6.1
66
4
19

314

6.0

106

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

188

TABUE 90
PEKCZNTAGBES OF SCATTSRSD kSTAPHASES
A. SHORT TRBAUCKNT
(Twenty p#rt# per v illi on)

8 B C a a 92
Hr#*
Soatt* Total
0
0
44
1
45
1
2
3
28
3
13
26
4
19
22
9
28
45
6
27
80
Total
Treated
60
193

>
0.0
8.8
10.7
50.0
66.2
62.2
74.1
41.4

SERIES 140
Soatt* Total
78
1
4
84
12
41
16
21
94
59
42
59
-

128

<
1.4
4.8
29.3
76.2
91.9
71.1

-

264

-

OUSBINB)
Soatt. Total
1
115
129
5
69
15
29
47
69
81
70
104
80
87
208

48.5

457

%
0.9
3.9
81.7
61.7
89.2
67.3
74.1
45.5

•
TABLE 51
FBRCS;TACaSS OF SCATTERED METAMIASES
A, SHORT TREATMENT
(Forty part# par m illion)

Hr#.
0
I
2
3
4
5
Total
TYaatad

SERIES 96
Soatt* Total
it
41
0.0
0
78
13.9
10
3.2
93
3
41.2
21
51
10
90.0
9
10 100.0
10
93

836

28.5

SERIES 120
Soatt* Total
%
0.0
0
48
73
1.4
1
12
18.9
65
33
19
97.6
98.3
13
12
4 ,100.0
4
48

188

OGUBINSD
Soatt. Total
It
89
0.0
0
7.6
11
145
9.9
198
15
47.6
84
40
23
91.3
21
14 100.0
14

25.5

Reproduced with permission o f the copyright o w n er Further reproduction prohibited without permission.

101

484

83.8

109
TABLE 52
PERCSNTACa» OF SCATTXRED klETAPRASES
A. SHORT TRKAHOENT
(Eighty parta par m illion)

ara.
0
1

2
3
4
5
6
Total
"Dpaatad

SBaSS 118
Soatt* Total
0
0*0
63
4
67
6.0
13
103
12.6
28
73
38.4
15
15 100.0
17
16
94.0
13 100.0
15
89

288

SERIES 122
Soatt* Total
_
Jt
0
0*0
75
9
61
14.7
18
111
16.2
13
86.7
15
12
85.7
14
4
4 100.0
-

30.9

56

205

27.3

OdCBIRED
Soatt* Total
%
0
138
0.0
13
128
10.2
31
214
14.5
41
88
46.6
27
95.1
29
20
95.2
21
13
13 100.0
145

493

89.4

TABLE 53
FSRCXNTAOBS OF SCATTERED KBTAHIASES
B. OONTINDOÜS TKEAIKENT
(Ona part par m illion)

Hra.
0
1

s
s
4

5
6

7
8
Total
Traatad

SXRISS 146
Soatt* Total _
62
1
1
73
47
2
7
43
52
54
72
84
58
67

^

_

1.6
1.4
4.3
16.3
65.4
75.0
86.6

•
-

156

354

»

44.1

SERIES 151
Soatt* Total
82
0
69
0
70
1
37
0
2
66
11
86
57
18
42
25
17
23
74

419

0.0
0.0
1.4
0.0
3.0
80.0
51.6
59.5
73.9
17.7

OOKBISED
Soatt. Total
1
144
142
1
117
3
80
7
1X8
36
127
65
76
124
42
25
17
23
830

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

773

0.7
0.7
8.6
8.8
30.5
51.2
61.5
59.5
73.9
89.5

160
TABLE 5 4
FERCEHTACaES OF SCATTERED KETAPHASE8
B. C0MTZ12D00S TREAWENT
(Flv# ports per milXlon)

Hre*
0
1
£
8
4
5
6
7
8
Totol
T^Peoted

SERIES 159
Seott* Totol
0
SO
1
65
18
57
17
36
39
55
49
SO

?(
0.0
1.6
51.6
47.2
70.9
98.0

-

-

-

-

-

-

-

-

-

124

261

47.5

SERIES 152A
Seott. Totol
%
0
57
0.0
0
77
0.0
62
1
1.6
12
72
16.7
22
57
58.6
48
09
81.5
77
70
90.9
52
56
92 9
72
76
94.7
277

556

51.7

aaUBJKKD
Scott. Totol
107
0
140
1
119
19
29
108
61
112
97
109
77
70
56
02
72
76
401

797

0.0
0.7
16.0
26.9
54.0
89.0
90.9
92 9
94.7
50.5

TABLE 50
FSRCENTAGSS OF SCATTERED KETAFHASES
B. COKTTKUOOS TREATlfSNT
(Twenty porte per m llllw )

Rre.

0
1
2
5
4
5
6
7
8
9
10
11
12
T otal
T reated

SERIES 128
Soatt* Totol
48
0
3
78
79
10
57
111
112
41
98
73
148
117

100

121

132
114
105
106
126

143
157
113
106
127

962

1373

;5
0.0
3.8
12.7
33.3
36.6
74.5
79.0
82.6
92.3
83.2
91.1
100.0
99.2
70.1

SERIES 154
Soatt. Totol
47
0.0
0
2.9
2
68
18.3
19
104
54.3
94
51
69.7
62
89
90
84.4
76
95.6
91
87
95.5
110
105
88 100.0
88
107
98.1
105
98.9
92
91
97.9
92
94
89
9A_ 97.8
86 7

1118

7 7.5

COMBINED
Soatt. Totol
0
95
è
146
185
29
205
88
201
103
149
188
239
204
251
205
231
220
219
244
194
205
200
198
218
.219.

5.4
15.9
40.9
51.2
79.3
85.4
88.7
95.2
89.8
94.6
99.0
98.6

2491

73.4

1829

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

0.0

Iftl
TABLE 56
PET’CEÎ’TAGBS o f sc a t t e r e d i^ETAPRASES
B . C C T m iU O tS TREATKENT

(Forty parts per m illionj

SERIES 95
0
1

z
5
4
5
6
7
e
9
10
11
12
Total
%aated

SERIES 125

1
1
15
18
49
77
44
101
121
121
116
105
-

54
59
62
87
83
96
81
111
126
122
119
109
-

2.9
1.7
24.2
20.7
59.0
80.2
54.3
91.0
96.0
99.2
97.5
96.3
-

0
4
25
49
94
101
106
87
72
109
127
105
125

59
96
87
105
128
108
114
90
77
no
133
107
129

768

1055

72.8

1004

1284

___2______
0.0
4.2
28.7
46.7
73.4
93.5
95.0
96.7
93.5
99.1
95.5
98.1
96.9
78.2

COLBINED
V ¥8 A
T
?«ax ____I
1
93
1.1
5
5.3
155
40
26.9
149
67
192
34.9
145
2n
67.8
178
204
87.3
150
76.9
195
188
201
93.5
193
205
95.1
230
232
99.1
252
96,4
243
210
97.2
216
125
129
96.9
1772

2339

75.8

TABLE 57
FERCEITAGEES OF SCATTERED KETAPHASES
B. CÜNTIWJ0U3 TREAINEWr
(Eighty parta par minion)

Hra.
0
1
2
3

4
5
6
7
8
Total
Traatad

SERIES 102
S oatt. Total
51
0
68
5
48
6
no
23
72
100
94
66
—
112
88
95
88
546

625

*
0.0
4.4
12.5
20.9
72.0
70.2
78.6
94.6
55.4

SERIES 115
Soatt. Total
47
0
2
99
96
5
27
106
55
n?
88
154
100
115
117
128
592

795

it
0.0
2.0
5.2
25.4
45.8
65.7
87.0
91.5
49.3

OOlCBIKED
Soatt. Total
0
98
167
5
144
U
216
50
217
125
94
66
134
88
227
188
221
205

0.0
3.0
7.6
23.2
57.6
70.2
65.7
82.8
92.8

1420

52.0

738

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

15»
TABU se
FBRCnVFAGBS 07 SCATTKHSD iXTAFHASBS
C. IKTStVITTBIT TREATMMT
(Forty parts psr B illion for 15 Blantss srsry two hours)

Hours
0
1
2

S

4
5
6

Soatt.
0
5
27
25
12
17
10

SERISS 126
Total
42
84
66
34
12
17
10

%
0.0
6.0
40.9
67.6
100.0
100.0
100.0

99
f

8
Total

9

9

100.0

103

252

54.2

TABLE 59
PKRCKNTAGBS OF SCATTSRSD MBTAFHASBS
C. TNTERLIITTSNT TRKATVKNT
(Forty parts psr BiUioa for 15 Blnutss svsry hour)

Hours
0
1
2

3

4
5
6
7
8
9
10
11
12
Total

Soatt.

SERIES 135
Total

0
11
42
55
76
80
88
88
83
99
lie
99
81

50
89
73
75
82
81
88
88
84
100
118
99
81

920

1056

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

%
0.0
12.4
57.5
75.3
92.7
98.8
100.C
100.0
98.8
99.0
100.0
100.0
100.0
87.1

168
TABU 60
PERCZKTAGXS OF SCAITBRBD &BTAPHA8E8
D, ÜM
TREATXD
(Root tip s oollsotsd a t oonssoutlTs hours)

Hrs.
0
1
2
3
4
5
6
7
8
9
10
11
12
Total

SERIES 12?
S oatt. Total
0
49
0
49
0
58
0
45
0
54
0
44
0
66
62
0
0
52
0
68
0
51
57
0
0
53
0

0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0

708

0.0

SERIES 132
Soatt. Total
1
64
7
70
2
61
1
58
7
84
4
59
3
63
15
63
7
85
0
59
69
8
-

-

%
1.6
10.0
32.8
1.7
8.8
6.8
4.8
23.8
8.2
0.0
11.6
-

5

51

9.8

60

786

7.6

OOt^BINED
S oatt. Total
1
113
7
119
2
119
1
103
7
138
4
103
3
129
15
125
7
137
0
127
6
120
0
57
5
104
60

TABU 61
PERGZffTACZES OF SCATTSRED RETAITL^SSS
D, inmULtTED
(Root tip s oollsotsd from asms onion a t asms hour)

Hrs.
0
1
2
3
4
5
6
7
8
9
10
11
12
Total

Root lo .

Soatt.

2
3
4
6
7
6
10
11
12
13
15
21

1
5
0
1
3
2
4
2
1
3
10
5

-

-

37

%

Total
94
128
124
99
111
105
109
81
111
125
105
163
-

10.6
3.9
0.0
10.1
2.7
1.9
3.7
2.5
0.9
2.4
9.5
3.1
-

1355

2.7

Reproduced with permission o f the copyright owner. Further reproduction prohibited without permission.

1494

0.9
5.9
1.7
1.0
5.1
3.9
2.3
12.0
5.1
0.0
6.7
0.0
4.8
4.0

164
TABLE 62
P3P.CEKTAGSS OF SCATTSRSD kSTAPHASES
TABLE 62A
ZERO HOQR OF 1SSATSD BULBS
SHORT TREATMENT
Parts psr m illion

Soatt•

Total

1
5
20
40
80

0
0
1
0
0

149
117
116
89
138

0.0
0.0
0.9
0.0
0.0

Total

1

609

0.2

TABIE 6£B
ZERO HOUR OF TREATED BULBS
OORTIKUOUS TREAT&SNT
Parts psr m illion

Soatt.

Total

1
5
20
40
80

1
0
0
1
0

144
107
95
93
98

0.7
0.0
0.0
10.7
0.0

Total

2

537

0.4

TABIE 620
SUtl’ARY OF XLL UhTRSATSD MATERIAL
Soatt•
0-Hour Short
0-Hour O ontlA U O us
ConseoutlTs Hours
Sams Hour
Total

Total

À

1
2
60
37

609
537
1494
1355

0 .2
0 .4
4 .0
2.7

100

3995

2.5

Reproduced with permission o f the copyright o w n er Further reproduction prohibited without permission.

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co

FERdrCiUSS or DISORCWUaaD AND TOTAL ANAPHASES
A. SHORT 1REAWENT

CD

C /)

(On# part par m lllioo)
8

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3
CD

3.

3
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S
■
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C

g
.
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3
■
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S

&
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C
%

<
(//>
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o'

3

SERIES 143
Total Total
Hour# Dlaorg. Ana. dit .
0
0
29
267
40
280
1
1
17
Z
1
254
S
34
1
248
0
4
32
251
0
13
230
5
0
16
253
6
T
1
11
269
8
16
1
234
32
9
z
281
10
0
27
254
U
0
26
273
1£
z
16
265
Total
Tr#at#d 9
280
3092

SERIES I47A
Total Total
Dlaorg. Ana. DIT.
0
236
18
0
13
214
212
1
14
28
220
1
244
19
1
21
269
1
201
0
11
14
260
4
0
230
12
0
31
240
0
235
14
17
222
0
220
16
1
9

210

2772

OCMINED
Total
Total $
i
dit . Ana.
Ana.
Diiorg.
Dltorg.
47
0
0.0
509 9.9
S3
1.9
1
494 10.7
466 6.7
2
31
6.5
2
3.2
62
468 13.2
2.0
1
SI
495 10.3
34
2.9
1
499 6.8
0
27
0.0
454 5.9
20.0
529 4.7
5
25
28
3.6
1
464 6.0
2
63
526 12.0
3.2
0.0
469 8.4
0
41
0
0.0
43
495 8.7
32
3
9.4
485 6.6
18

490

3.7

5864 8.4

8

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TABU 69
FERCEtTTACaES OF DISORGANIZED AND TOTAL AKAFfUSSS
B . OONTINUODB IM A M N r

8

TD

(Plv# part# p»r m illion)
CD

CD

CD

"O
O
Q .

O
3
■
D
O
&
O

c

%

(/)
o'
3

Bouri
0
1
2
3
4
5
6
7
8
Total
%aat#&

SKRHS 139
Total Total
Disorg* Ana. DIt *
1
1
4
S
8
1

31
14
45
22
19
16

-

-

-

-

-

-

12

114

267
256
279
230
215
213
-

1193

SBIES iseA
Total Total
Oisorg. Ana. DIt .

GGKBI
Total
Total
a
Ditorg. Ana. Dlaorg. DIt . Ana.

0
0
0
0
2
3
5
3
6

41
27
34
19
21
28
21
15
20

257
240
311
300
273
257
256
225
222

1
1
4
3
5
4
5
3
6

72
41
77
41
40
44
21
15
20

1.4
2.4
5.2
7.3
8.0
9.1
23.6
20.0
50.0

524
496
590
550
488
470
256
225
222

13.7
8.3
13.0
7.7
8.2
9.4
8.2
6.7
9.0

19

165

2064

31

299

10.4

3277

9.1

s

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FER C Xm C SS o r DISORCUKIZED and t o t a l P d lk m s s s
B . OOSnNDODS n S A 3 1 i» T

(/)
C
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(Twenty parte par m illion)
8

CD

C

3.

3"
CD

CD

■
D
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C

g.
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D
O
&
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%

SE Sie 128
Total Total
Houra Diaorg* Ana« dit .
0
28
230
0
232
ù
14
1
292
z
0
6
9
0
3
228
4
4
236
9
2
10
5
221
4
243
6
6
7
6
10
236
8
6
6
259
2
9
3
245
12
216
10
14
222
11
8
9
12
0
0
219
Total
92
2769
Traatad 46

134
Total Total
Dlaorg. Ana. dit .

OGMBINKD

sbrus

Total $
Total
i
Dlaorg. Ana. Dlaorg. dit . Ana.

0
1
0
2
4
9
5
4
6
8
5
4
3

21
12
8
14
12
16
10
11
6
10
10
9
6

213
220
268
276
256
277
272
275
253
245
256
248
242

0
1
0
2
8
11
9
12
12
10
17
12
3

49
26
16
17
21
26
16
21
12
13
24
18
6

0.0
3.9
0.0
11.8
38.1
42.3
56.3
57.1
100.0
76.9
70.8
66.7
50.0

51

124

3088

97

216

49.9

443 U .l
432 5.8
500 3.2
504 5.4
492 4.3
498 5.2
515 3.1
511 4.1
492 2.4
490 2.7
472 5.1
470 3.8
461 1.3
5857

3.7

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XT4
TABLE 7 3
FBRCZNTAGXS 07 DISORQAKIZED AKD TOTAL ANAPRASSS
0. INTRKITISNT TRSAIlEXirr
(Forty port# p#r m illion for 15 minute# every two hours)
SERIES 126

.

Hour#
0
1
2
3
4
5
6
?
8
Total
%eated

Dlaorg.
1
0
10
3
6
5
5
»
2
31

Total
Ana.
31
15
35
49
20
7
5

Dlsorg.
3.2
0.0
28.6
6.1
30.0
71.4
100.0

—

Total
Div.
227
228
218
227
211
184
206

—

—

Ana.
13.7
6.6
16.1
21.6
9.5
3.8
2.4
_

2

100.0

78

2.6

133

23.3

1352

9.8

TABLE 74
PBROBOTAGKS OF DISORGANIZED AND TOTAL ANAPHASES
0. INTERMITTENT IREATKENT
(Forty pert# per m illion for 15 minutes every hour)
SERIES 135
Hour#
0
1
2
3
4
5
6
7
8
9
10
11
12
Total
"Rreated

Dlsorg.
0
3
6
5
8
12
6
16
15
12
8
3
6
100

Total
Ana.
16
12
51
30
24
39
9
20
18
14
8
3
6

Dlsorg.
0.0
25.0
19.4
16.7
53.3
30.8
66.7
80.0
83.3
85.7
100.0
100.0
100.0

214

46.7

Total
Dlv.
228
264
235
257
257
273
234
247
233
256
256
245
244

Ana.
7.0
4.5
12.3
11.7
9.3
14.3
2.8
8.1
7.7
5.5
3.1
1.2
2.4

3019

7.1

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

CD

"O
O
a
c
g
a.
TABUï 75
■
D
CD

PSRCSmCSS OF DIS0RaA.M2aSD AI3 TOTAL AIIAPKASES
D. ONTOÏATBD MATERIAL

C /)

o'
3

(Root tip s eollseted t t consecutive hours)
CD

8D
■
C
5IT
3
CD

C

3.

3"
CD

CD

"O
3

Q .

C

a
O

3

TD

O

CD

a
O
■
D
c

CD

Hours

SKIES 127
Total Total
Dlsorg. Ana. Div,

SERIES 132
Total Total
Disorg. Ana, Div.

0
1
2
3
4
5
6
7
8
9
10
11
12

0
1
0
0
0
0
0
0
0
0
0
0
0

22
36
22
33
26
46
28
24
26
25
51
21
34

214
235
229
256
231
237
261
221
223
240
229
227
243

2
10
4
11
3
1
3
6
1
0
3
0

13

Total

1

576

3046

44

345

24
49
29
24
40
26
21
25
39
28
27

-

Cd'BIKEO
Total
i
Total i
Disorg. Ana. Disorg. Div. Ana

157

2
11
4
11
3
1
3
6
1
0
3
0
0

46
85
51
57
66
72
49
49
67
55
58
21
47

4.4
12.9
7.8
19.3
4.5
1.4
6.1
12.2
1.5
0.0
5.2
0.0
0,0

2861

45

721

6.2

251
272
240
230
258
212
240
241
296
232
230
-

465
507
469
486
489
449
501
462
521
472
459
227
400

9.9
16.8
10.9
11.7
13.5
16.0
9.8
10.6
12.9
11.2
12.6
9.3
11.8

C /)

W

o'

3

5907 12.2

175

TUSIX 76
FERCIEKTAOBS 07 DISCRQANIZZ) AKD TOTAL AKAPKASBS
D. UNTREATED kATKRIAL
(Root tip# oolloetod from ##m# onion nt nmm# hour)

Root No*

Z
3
4
6
7
8
10
11
12
13
15
21
Total

Dlaorg*
Ana*

Total
Ana.

%
Dlaorg*

0
8
1
1
2
2
0
0
0
1
9
0

34
63
37
31
48
48
29
26
51
43
48
47

0.0
12.7
2.7
3.2
4.2
4.2
0.0
0.0
0.0
2.3
18.7
0.0

24

500

4.7

Total
Dlv*

$
Ana*

454
7.8
498 12,7
413
9.0
422
7.3
448 10.7
450 10.7
6.8
429
429
6.1
442 11.5
459
9.4
441 10.9
491
9.6
5356

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

9.4

1T7

TABLE 77
FERCS^TAOES OT DISORGANIZED AND TOTAL ANAPHASES
TABUK 77A
ZERO HOURS OF TREATED BULBS

PmrtB per m illion
1
5
20
40
80
Total

Dlsorg*
Ana*

Total
Ana.

T

'

"■

Disorg.

0
0
7
5
1

47
57
48
53
51

0.0
0.0
14.6
9,4
2.0

13

256

5.1

TABLE 77B
ZERO HOURS OF TREATED BULBS
CONTTNUOUB TREATMENT
P arts per m illion
1
5
20
40
60
Total

Dlaorg*
Ana.

Total
Ana.

%
Disorg.

1
1
0
5
S

50
72
49
51
SO

2.0
1.4
0.0
9*8
4.0

9

272

3*3

TABIE 770
SOKKARY OF ALL UNTREATED MATERIAL
Total
Total
Disorg.
Ana.
Dlsorg.
0—
Hour Short
0-Hour Cont inuous
ConseeutiTe Hours
Same Hour

13
9
45
24

256
272
721
505

5.1
3.3
6.2
4.7

Total

91

1754

5.2

Reproduced w # permission o f ,he copyright owner Further reproduction prohibited wtthout permission^

178
TABLE 7 8
PKRCKUTAGES OF CELLS CONTAINING "REDUCTIONAL"
OR **SBaRSQATIOMAL" GROOFINQS
A. SHORT TREATMEIfr
(One pert per DllXlcm)

Hours
0
1
2
5
4
5
6
7
8
9
10
11
12
Total
T r e a te d

SERIES 143
See*
Total
3
176
2
181
5
213
5
183
3
183
1
196
216
3
4
234
5
203
228
4
8
198
2
214
197
2
42

2442

SERIES 147A
See*
1
0
2
0
1
2
0
3
0
1
1
1
4
18

Total
193
188
167
147
169
214
186
207
180
187
180
172
181
211?

See*
4
2
5
3
4
5
3
9
8
8
9
5
6

OORBINED
Total
369
336
380
332
382
410
374
441
383
412
378
386
378

87

4599

%
1.1
0.6
1.3
0.9
1.1
0*7
0*8
2.0
1.3
1*2
2.4
0.8
1.6
1.3

TABIE 79
PKHCEKTACTS OF <3EL IS CONTAINING "REDUCTIONAL"
OR "SECREGATIONAL" OROCFINOS
A. SHORT TRSAmSNT

(Five parts per m illion)

Hours
0
1
2
3
4
5

6

7
8
Total
I V e a te d

SERIES 138
Total
See*
208
1
1^9
5
7
228
234
1
183
6
100
1

—

a.
20

944

tiZëltlES 182B
Total
S ee*
180
2
173
1
198
9
223
4
182
6
215
3
172
3
206
1
41
2

29

1410

Sag*
3
6
16
8
12
4
3
1

s

CORBIND)
Total
388
37i
426
457
368
315
172
206
Ü . ..

49

2354

Reproduced with permission o f the copyright owner. Further reproduction prohibited without permission.

0.8
1.&
3.8
1.1
3.3
1.3
1.7
0.8
2.1

m

TABIX 8 0
FERCSNTACBS OF 0ELL8 OONT4IMIMO "RXDDCTTONAL**
OR **SBaRXaATIONAL" OROUPINQ6
A. SHORT IRXAHCENT
(Twenty perte per m illion)

92
Houre
0
1
2
3

4
5
6
Total
"Dpeeted

Seg«
7
8
5
4
8
5
4
30

Total
157
187
154
140
154
191
121
947

SERIES 140
Sag*
Total
8
200
7
827
9
205
206
1
3
222
157
0
-

-

20

1017

nrwpjvgn
Sag.
15
15
14
5
9
3
4

Total
357
414
359
346
376
348
121

$
4.2
3.6
3.9
1.4
2.4
0.9
3.3

1964

2.6

50

TABLE 81
PBHCKNTAGBS OF CELLS COETAIKING "REDUGTIONAL"
OR "SEŒŒQATIONAL" GROÜFÏKQS
A. SHORT TREAWWT
(Forty parts per m illion)

Seg.
Koore
0
4
“T----------- 18
2
4
3
16
4
8
5
5
Total
Throated 51

98
Total
101
197
198
165
159
58
777

SERIES 180
Sag.
Total
4
174
185
8
174
16
6
163
121
1
0
25
31

668

COMBINED
Seg.
8
26
20
22
9
5
82

Reproduced with permission o f the copyright owner. Further reproduction prohibited without permission.

%

Total
325
382
372
328
280
83

2.5
6.8
5.4
6.7
3.2
6.0

1445

5.7

IBO
ThBlM 8S
ITOOSimiaES OF CSLLS CORTHIHIIIO ""REDDCTIQNÀL"
OR "SBGRXQATIOIUL'* OROOPIRQ8
A. SHORT TREAwmrr
(Eighty p arts par B illion)

Houra
0
1
2
S
4
9
6
Total
Traatad

SERIES 1X8
Total

SKIES 122
Sag.
Total

9
9
10
11
2
0
4

199
194
299
197
151
141
177

10
9
7
2
1
2
-

229
207
273
141
193
151

96

1113

17

909

-

Sag.

Total

%

19
14
17
19
9
2
4

424
401
526
998
904
272
177

4.8
9.8
9.2
9.9
1.0
0.7
2.3

59

2018

2.6

TABLE 83
FERCENTAOBS OF CELLS OOMTAliaNQ "RZDDCTIuNAl"
OH "SEGKEGATIONAL'* GROUPINGS
B. CONTINUODS WEATkENT
(Om» part par m illion)

Hours
Sag.
0
1
2
1
2
2
S
4
4
0
5
0
6
2
7
*
e
Total
IVaatad 10

CWBINED

SERIES 151

146
Total
179
219
213
189
169
198
186
»
—

1170

Sag.
S
1
9
0
11
18
9
1
2
36

Total
188
182
229
172
227
258
180
177
128
1999

Sag.
4
9
9
4
11
15
9
1
2
46

Reproduced with permission o f the copyright o w n er Further reproduction prohibited without permission^

%

Total
967
401
442
997
396
456
966
177
128

1.1
0.8
1.1
1.1
2.8
5.9
1.4
0.6
1.6

2729

1.7

m
TABIX 8 4
PTOCENTAOES OF CKLIS CONTAIIONO "RKDOCTIONAL"
OR "SECaaGATIOKAL** OROOPINOS
B, OONTIKUODS TOBAIWCNT
(Five parte per m illion)
tiJSKiSS 139
Heure
0
1
2
3
4
5
6
7
8
Total
Treated

8eg.

SERIES 152A

Total

Seg.

COMBINED

Total

Seg.

Total

%

3
4
9
2
7
3
-

191
209
219
190
184
186
-

3
5
3
6
4
20
6
2
12

184
196
257
255
232
220
224
199
185

6
9
12
8
11
23
6
2
12

375
407
472
445
416
406
224
199
185

1.6
2.2
2.5
1.8
2.6
5.7
2.7
1.0
6.5

25

984

58

1770

83

2754

5.0

TABLE 85
FEROSNTACaCS OF CELLS COKT41A11JO **RSDDCTIONAL**
OR "SSCaRBOATIGNAL** GROUFINOS
B. OONTIHUODS TREATMENT
(Twentj parts per m illion)

Houre
0
1
2
3
4
5
6
7
8
9
10
11
12
Total
Treated

SERIES 128
Total
Sag.
2
153
210
8
221
0
223
4
226
4
209
4
236
2
224
1
230
1
240
2
200
6
209
5
215
5
42

2643

SERIES 134
Total
Sag.
159
0
266
4
287
19
25?
16
241
14
257
4
2
257
8
256
10
235
8
233
7
245
2
234
232
3
97

2904

Sef.
2
lA
19
20
18
8
4
9
11
10
13
7
8

COMBINED
Total
3lÇ
416
478
480
467
466
493
480
465
473
448
443
447

139

Reproduced with permission o f the copyright owner. Further reproduction prohibited wi
without permission.

5547

$
0.6
é.9
4.0
4.2
5.9
1.7
0.8
1.9
2.4
2.1
2.9
X»6
1.8
Ca6

la t
TABLE 8 6
FKROENEAGBS OF OXLLS OONTAIMIKQ "RXDOCTIOKAL**
OR ’♦SEGKEOATIONAL" GROOFINaS
B. CONTINUOUS HQBAIUXNT
(Forty ports per m illion)

m ss
See*
5
29
21
11
20
24
9
14
7
18
1
1

Hours
0
1
8
S
4
5
6
7
8
9
10
11
IE
Total
TTsated 151

99
Total
162
190
198
191
194
199
182
200
212
179
209
174
-

2116

SRIES 129
Seg.
Total
0
196
9
223
4
197
4
212
11
228
9
216
9
210
4
199
6
192
10
219
7
231
7
229
221
9
81

2573

COMBINED
Seg.
9
30
29
19
31
29
18
18
13
28
8
8
9

Total
998
418
399
403
422
411
392
399
404
394
436
403
221

1.4
7.3
6.3
3.7
7.4
7.1
4.6
4.6
3.2
7.1
1.8
2.0
4.1

4331

9.4

232

TABIE 87
FEl^CENTACSES OF CELLS CONTAININO «RXDOCTIONAL**
.
OR "SEGREQATIONAL" GROUFENOB
B. CONTINUOUS TREATMENT
(ElgHty p arts per m illion)

Hours
0
1

2

SERIES 102
Total
Seg.
5
4
6
10
30
18

3
4
5
6
7
19
17
8
Total
Treated 104

SERIES 119
Total
Seg.
166
1
236
9
212
2
266
21
254
10

Seg.

207
199

29
19
16

232
208
222

6
9
8
31
40
18
29
34
33

1423

94

1630

198

144
199
204
227
227
204
—

•

-

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

COMBINED
Total
310
431
416
493
481
204
232
419
381

$
1.9
2.1
1.9
6.3
8.3
8.8
10.8
8.2
8.7

3094

6.4

las
TABUC 8 8
PKHCXNTAOES OF CELLS CONTAINIKO "REDOGTIONAL*
Oft "SlQRECUlTIOmL" CROUPINOS
C* tJNTftSimD
(Root tip# ooll#ot#d #t

Hours
0
1
2
S
4
5
6
7
8
9
10
11
12
Total

SERIES 127
Sag.
Total
1
157
2
168
1
181
3
186
0
156
4
142
0
189
0
172
0
169
1
183
0
158
1
158
0
171
13

2184

oonaeoutlT#

SERIES 232
Sag.
Total
1
186
7
182
2
170
5
178
1
196
3
157
2
188
5
186
0
201
1
165
1
160
-

Sag.
8
9
3
8
1
7
8
3
0
2
1
1
0

-

0

128

26

8091

hours)
COMBINED
Total
343
344
351
364
358
899
377
358
370
348
318
158
293

39

4275

TABLE 89
FEKCSJ'.TAOES CF CELLS CONTAINIIIO •REDOCTIONAL"
OR "SBGEtSQATIONAL** GROUPINGS
0. UNTREATED
(Root tip s eollsotsd fran ssms oaioa s t ssme hour)

Hours
0
1
2
3
4
5
6
7
8
9
10
11
12
Total

Root No.
8
3
4
6
7
8
10
11
18
13
15
21

SBtISS 137B
Sag.
Total
330
3
6
366
335
6
330
4
348
4
0
334
4
341
335
8
0
306
10
345
12
385
376
1
-

-

58

4071

0.9
1.6
1.8
1.8
1.8
0.0
1.2
0.7
0.0
2.9
3.7
0.3
-

1.3

Reproduced with permission o f the copyright o w n er Further reproduotion prohibited without permission.

%
0.6
0.3
0.9
2.2
0.3
2.3
0.5
0.8
0.0
0.6
0.3
0.6
0.0
0.9

IM
TABLE 90
EERCEKTACTS OF CELLS COWTAININO **FŒDUOTIONAL"
OR *»SBORE(UTIONAL" CROUPmOS
TABLE 9QA
ZHIO HOURS OF TREATED BULBS
SHORT TREATKENT
P#rt# per m illion

Seg.

Total

1
5
80
40
80

4
3
13
8
19

369
355
357
385
484

Total

49

1830

%
1.1
0.8
4.8
2.5
4.5
2.7

TABLE 90B
ZERO HOURS OF TREATED BULBS
CONTINUOUS THBA'aaOT
Parts per m illion
1
5
80
40
80
Total

Seg.

Total

%

4
6
2
5
6

367
375
312
358
310

1.1
1.6
0.6
1.4
1.9

23

1728

1.3

TABLE 900
SaOüfflT OF ALL UNTREATED UATfillAL
Seg.
0-Hour Short
0-Hour Continuous
ConseeutiTs Hours
Same Hour
Total

Total

%

49
83
39
58

1830
1722
4275
4071

2.7
1.3
0.9
1.3

163

11898

1.4

Reproduced with permission o f the copyright owner. Further reproduction prohibited without permission.