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t
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induced Polyploid PeriOliiiisX CMmeras
,Itt Solanum Tvbwto&m
■ iby
Eiehard ilwln lake**'
A THKSI3
Presented to the Faculty of
The Graduate College in the Bbieereity of Hebraeka
m Partial Fulfillment of Hequtrements
■ For the ^Degree of Pli* »*
Oepertoent of Botany
Lincoln, Nebraska
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UMI Number: DP13661
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induced polyploid, Perleliaal QhiMvu In Bolnmm Tx$>®mem
XXMXM
' EX0BA1P BAKBR
3H9BB ESI smstTXBSOV or MBWUtM PSOfSBSOR If 1* W&SKBft
fhi® Investigation m ® begun la fell# fall of 1988 during the
initial period (4S) «f enthusiasm for the production ©f pdly-
plold plant®.by ©olcMclme treatment* fb® principal objective
was the det#*iM»abi©» ©f bb® inherent vain# of an Increase la
the cbromoaas© number of Solteus* for oiMnoooiol potato
to
production, this end aeeda and seedling® of aa inbred at rain
of potato were treated with ©olcMcin#* The treatmenta resulted
la the production ©f polyploid, perieliaal chimeras which fail
ed to develop flower®* Morphological, anatomical, and cyfco-
logical studies of these plant® form the basis for the entire
discussion in feM® .paper*
MVtWS.
The. developaeat of polyploids and perlclinal chimeras re
presents two separate fields of Investigation with distinctly
different contributions* The- indtwtioa of polyploidy in plant®
has made 'possible the production of anpMdlploi&a,' fertile inter*
specifie hybrid®, and new forms (10).* Studies of periclinal
chiaera® have contributed much toward a better understanding of
developmental 'morphology (18)* ' The close relationship of each
of these 'two .iaportant field® of Investigation to the present
discus®ion makes it worth while to review briefly some of the
literature concerning them*
nrl',r'' fhe author wlshear't© Express M s sincere appreciation to
Associate Professor I* E* Walker of the Department of Botany
under whose supervision this investigation was carried out, to
Professor h* 0* Wemer of the Department of Horticulture for
supplying the Material used, and to the Departments of : Horti
culture ted Agronomy for furnishing equipment.
398987 - ; r
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s
Polyploids b##» obtained in many species of Solanxjm
by the deeapitation-callus method, bisdotroai and loos {30}
obtained a diploid tomato plant fa?m a haploid of spontaneous
origin* When this same technique was tried on tbs diploid a
tsbraplold was produoed* bitroganling the possibility of
mutations th® in, to# and to plants oonslsted of different
multiples of the same chromosome set#, Hoguchl (34) also
oamS'Od the 'formation of ■ batraploid tomato plants, y©rg«»*#»
{19, 20) and Xftgftv* (22) reported tbs- successful us© of this
method for ether, species of Solanum*
Shortly after the introduction of. the ©olchtoin# taebatfu®
yobastoa# (17) treated 276 tubers of £# tuberosum with eel*
chicineianolin paste and reported the production of .one plant
vhloh had 96 in the- root tip# of the treated
generation, fbe treatment of 1160 seed* of three seedlings of
the above speslea resulted in one plant which had 9# somatic
OfaremuKMiss in the root tips after it had been propagated
through one aaesaiel generation by cuttings or -tubers# An
im&mmm in the number ■ of somatic toroMNHMM in. epidermal
cell#' ©f some treated plants was indicated by at©»atal measure-
meats#- fhis work demonstrates elearly the difficulty ©f ob
taining a doubling of; the chromosome nutoer In £# tuberosum.
Tim untreated plants were vigorous and bloomed profusely toil#
the treated plants were -swell and failed to produce flowers •
twbaijii and Runquist ($B) treated seed of j|* tuberostM#
variety triumph# with ©©ichicifi# -and v*po*t*4l the production
of two plants with 96 chromosome# In the root tips* fh#
morphology -and detailed cytology of these plants was not given*
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$
Tim
somatic chromosome number has been doubled in other
species of Sojatam fear colchicine 'treatment"* Johnstone (17) ■
secured a ' doubling in S* jamcall, JS* tinman, S* bulbooast-*
anum* S* andigenum, and: s. neoantipoviohll.
I^rieliasl'V&ribeeree of'*w different type# have been
reported'in the literature* Satesen (§) and Clausen and
Soodepeed (7) described perlclinal chimeras of natural no*
eureno® in nicotians tab&oua* 'She epidermis differed from
the .internal tissues* Asexual propagation of the internal
constituent was accomplished by the use of root cuttings*
Aaaejeva (1) used an ingenious method to demonstrate the per*
iclinal nature of many ekt»ers# caused' by bud mutations in
ISO varieties of £* tuberosum the eyes of the tubers were
partially cut out or scraped to stimulate adventitious bud
growth from the internal tissues* the epidermis and inner
tissues were found to be involved'either alone or in combina
tion in the various mutations* fhe relationship of this
work to the present is treated in the discussion* Crane (8),
.Momtosh (33), and Clark (5, 6) used AsseJ#va*s method to
study bud mutation# in & m m m M varieties of £* tuberosum*
Jorgensen (80) reported a polyploid, perlclinal chimera of
!* whieh originated fro® a decapltat-ed^c-allus treat*
meat* fhe -outer two or more layer# were stated to be.tetra*
plold, and the internal tissue# diploid* A precise dec*
arlptlen was not given*', irenke CM) and iange (25) produced
perieliael chimera# by decapitation at the point of graft,
union between two species of Solanum. Winkler (46, 47* 48)
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4
who 'originated ib® term chimera, uee&'tMs method with jS*
jyeoperelem and £* nigrum* JorgeaM'and;-Oran*., (fil) reported
on .graft. hybrid* involving §# lyeeeereiena# S* slsyiabrlfollum.
\yfam*
g;«. f* nigrum w . graolle. and £* ISigSSg** Winkler
(48) ant. Lange (ft)’ were able t* propagate some of their plant*
by steOMinl methods* Satina, Slake*!®* #. ■ and ivory (17) obtained
polyploid, periclin&l ohtaorai by ooloMoine treataeat of
.Befog* Tin*©® IMependent gown layer# were reported
for the. shoot apex* Many . of in, 4n# sad 8n
layers were deaoaetreted* The relationship of eeeh of these
layer*, to the development of the shoot was dleeueeed# .Satina
end Sisk**!** (56) reported the enlede&ee of three gem layer*
In the floral apex* They deeeribed the relationship of each
to the develepeeut of the 'leaf, sepal, petal, and stamen*
-defer* beginning a dissuasion ef the present investlga»
tlon It I* necessary to define pertain tense* solanun tdbop*
osum. which has 48 aematle chromosomes, has ocramonly been
ref owed to a* a diploid* However, it has'been considered
possible' that" the basle chromosome number may be six or
twelve.
':2*wr«tte* (§7) studied the secondary association of ©hram-
©semes and suggested that the basic h&ploid number in. Solanum
1* six* Merer, the majority of the- evldenee fixe*, the baeie
number at twelve* fropaeh (£0) studied meloaii in apeele*
with £4, 36, 40, and 78 somatic ehromoaomes and found that
etkremeaewe groups ef high** valencies were more frSfueht in
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5
polyploids# S®\ suggested- that secondary association
of ohroMOOOBMO- should be-taken us significant evident# of
homology only when primary ass ©elation* ■ of similar valency
sis© occur and eoneludod that the-bail# chromosome number of
tii® toot ion fafee.yi^lm is' prbfeably too loo* Lunden (SI)
decided from genetic data that j|* tuberosum is an autototfop*
loid# The work of' %mm. .and lid# {&) m ■ the inheritance of
reset ion to oowun scab in tb* potato substantiates l»ua#®a«*
hypothesis* this Mono that a tstr&plold has 48-Muafeld
chromosomes and an @©t©pl®id has ©§*/All of tfao dloooooloa ■
which follows is based .m this viewpoint *
'wmaxtm wtmm
aid
The istered vtMUi of j|* tdberoita* which was. employed .in
this study Is one whleh is msed in tbs potato breeding pro
gram ©f tbs Department. of' BortiouXtnro at the University of
ibbraska and is designated number Aid (Sinn® sot a number
4X*1*1*7«M.)« It was ehosan primarily because it produces
an abundance of viable seed- when grown in the greenhouse
under 1? ©r 24 hours of light per day -and at temperatures
between id and 70* fy Also it is sufficiently hsnosygotti
t©,insure- a .rather uniform population ©f seedlings# All
plants used In this experhwsst were grown in the greenhouse •
Soil was used as a growth medium la all of tfaie work* Seed*
tr'bt&ers ©f untreated 'and treated plant* were started dur
ing September of each year from 12SMJ41* tuber® war®
planted ■ directly in fiw»iaeh pots# but:v seeds were planted
in flats and the resultant seedlings were shifted -to five*
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$
tmh pots* It was possible- to-produce **tl*faetory arterial
for ©ytologioiiX m snatoaieal study continuously until tb®
first ©f* second week of May# Davlroxuatntsl. conditions were
Maintained as described above in an a&teapt- to promote f%mm
prbtuotion 14, 44} during the. first'.SIO day# of. tbs greaafaeaae
seaeon* ' Bight and twelve .hour light periods: ware also used
for so** of fcht treated plants during the last yew?* During
the last 50 days of aaeh' gveeathoiiee season snrlroansntsl con
ditions -wort adjusted, for tuber production (43)*
•■ During the first . m m m dry seeds wen# placed cm sterile
filter paper.la:at# rile petri.dishee and .were then yetted with
the following concentrations of sgnsens solutims ©f ©oIsMeinei
*01, *03, *1, *f, :lg i, 5, 4,' sad 1- per-cent* The periods
©f exposure were f4,-48, ft, and 96 hour® at 70° F. for each
eeneeotration* ebeelr samples were wetted with distilled
water f©r identical intervals* Twenty«*flv# seeds were used
for each of the OS possible ©©aceatraiioii^itste combination®,
treated seeds were rinsed in distilled water at the ©ad of
the exposure interval#
la IfOO two additional types of treataenfc were used* ■
me consisted of-gsradnetlag seeds- ©a solst soli until the
radicle was barely e»ergin#* The seeds were then transferred
to petri dishes o«nt*lning ©lean sand which was wetted with
a #J per seat atueous solution of aa&chtel&ft* The following
exposure Intervals were uoodt M* 48# and 71 hours* too
hundred seeds were exposed to each of the three- possible
eonoentration»tia» emfeinaftloiuu The seeds -were- then
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