INHERSTANCE OF SPEED OF GERMINATION
EN SMGAR REETS
(BEA VULGAMS‘ L.)

Thesis far ”10 chmo 0? pk. D.
MICHIGAN STATE UNIVERSETY

Thomas E. Sedlmayr
1960

This is to certify that the
thesis entitled

Ismxuscs or am or Gsmmmxos In
em BEETS (am vummxs, 1..)

presented by

rho-is B. Bedhsyr

has been accepted towards fulfillment
of the requirements for

Phe De degree in Fm Crop!

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filminé S .Setflvaqyr
candidate for the desree (.

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Final eveminetion: February 2;, 1903

Inheri*avce of Speed of (erminafien if in er

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Cutiine cf
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Kinor subjects: Geretice, Path. Juveistice

Hinrrayhicai Items:

Hiwtl Schocd : Bermaijctizfie, iknwron, }Njn‘aljf, lfik‘

Undergraduate Studies: Cnivergitf OL niTiCULVKTC,
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Exocriehce: Plant areefler (corn), Agricuiiurei JQFQfirCh

InstiLute of the hungariun Acader

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.—

he seeds* of sugar beets possess unsatisfactory germination
characteristics from the agronomic point of view. Thus it is not too
surprisinr that 5—10 percent of the planted sugar beet acres;e is lost
annually, largely because of irregular or poor seedling emergence
(Agricultural Statistics, 1953), and many acres have to be replanted

(COR 19u7, thHCL 1955). Unsatisfactor; gerwination and slow field

[11

emergence also account for beet stands considerably thinner than the
optimum, causing reduced yield and sucrose content (sax h» lead, nlCnvL
1955). Although rapid emergence may not be the crucial factor every
year, in some years and in certain fields it may become one of the hOSt
important factors determining yield.

With the increasing use of monogerm and polyploid su ar beet
varieties, the demand for hiih quality seed is becoming extremely urgent.
Seeds of polynloid sugar beet varieties generally have a lower germina-

tion capacity and a slower rate of germination than do the equivalent

 

*The term sugar beet "seed" will refer to either monogerm or multigerm
fruits. For multigerm fruits the term ”seedaall" will be used, for
monogerm fruits the tenn "fruit". The sugar beet seed, as defined in
the botanical terminoloyy (frequently referred as to "genn" in the
sugar beet literature), will be consequently designated as "true seed"
to avoid possible confusion with fruits and seednal s.

diploids (PETO and hlLL l9h2, )tJlerd 19)?). To fulfill their intended
puroose , i. e. snace planting to a herfect st and, seeds of mono erm vari—
eties reo uire be tte er germin ati on ctarrcteristics than do multigerm
varieties. With mono;ern seeds, a high oercent of germination COHUinCd
with rapid and uniform ever ence is a virtual necessity.

Rabid enerwence is especially in ortaht vlere low temnerature,
poor aeration, critically low soil moisture, or soil crusting create
temporarily unfavorawle environ ental conditions. Slow emeraence may
delay prompt cultural operations, cauiing disadvantageous comyetition
between the super beet seedlinss and weeds. The -everity of certair
diseases mat al:3o be consideraoly decreased by rayid energence an.
vigorous subsequent vlowth (AFANASIEV JQAZ). 3 iurce a dire ct correlali3n
exist 3 between the length of growing season and yield and also sucrose
content(hUChCL319L6), earlier energence rroviies a lonier period of
growth for beets, and consequently a higher sugar yield.

The agronomic value of the sugar beet seed has been determined
largely on the basis of the germination capacity (oercent of seedhails
producing at least one seedlin3fl /, or on the oasis of the total number
of seedlines per 100 seedballs, under certain laboratory conditions.
Recently, however, more and more enohasis has been flaced on rate and

vi or of termination (QLATLd 1951, Silhjl 1955).

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A8 a most obvious aggroach, dit‘ere
treatments of the seedtall have been used 0 accelerate geruiratgon in

surar beets (H LTER 1951, DERTER and IlVa C”" 1W 9). However, verv

,JJ

little has been done to imrrcve s;'eer_ of gerxination or ether seed

 

 

 

 

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characterist V; 9
sequence of the gener
teristics of the suga
larzely ov environten
temperature, etc.).
The main pur;

~35 s . . n . . ‘ \
oi germination in at;

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ant breedin, ass of . 33.Ls tan a is lost cer—

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A] 1.2! ’9 L309; Lift"), ad 1' UJUT ~3 l‘Jn ' r104“ : elf...» {lullal’l Cii'PrdC"

. o '1 o
r beet seed are innately poor and are ini3uenced

fl.

tal “actors (see bail maturity, stora e conditions,

(vs

V

nose of this thesis is to determine whether speed

\ o u ‘ ‘ J‘ ‘ r“ ‘ .- ’ 4"
’ar beets is renetically controlled and to u.:v

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3

_n imorovina ttis character?

ann‘

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A number of factors are thou ht to we resoonsicle for
or slow emergence of sugar beet seeds. These factors can be classi. ied
conveniently into two main grouns: (l) l'n_ ernal factcrs, inherent in
the sufiar beet seed ("eecc"ll or true seed) itself, and (a) External
environmental factors, affecting the emerjence of the planted su;ar
beet seed under fEEld conditions.

Fumerous ercernal factors, such as soil moisture, soil tomfPT3~
ture, aeration, soi structure and compaction, tillage practices, various
seedbed preferation tech 1ruh, fertilizer anrlication, cron rotation,

etc. have irnortant roles in obtaininj earl; and uniforn energvz as

(u

(3

(L'
\-

have been shovn by AidhlfiiLD (1951), ha;V¢V (1959), thfuZ (JQSl), Hlllim
and DTYTsa (1951),L . Ta? and stitugcm (1999), STcUT, et al. (1957),
hOFFOgfl (1953), uossm‘ D and DHATLi (195;). Be caus e of the scope of ttis
thesis, we vill li it tho det"i led review and discussion, however, solely
to the internal factors
Environmental condi ions durin; tle devc lourent ano rinsrin, of

eeds can have a Creat influence on the subsequent gerxir;tjon and

owth of the re :ul inf flant (317D and 3&5? l9l3). In suiar beet ,
F“‘WJLTCN (l95h) found that under field conditions phosphate and reavy

nitroyen fertilizer a olication increased the germination caoacity or

the seed by 8—10 nercent. titro en alone had a less firehounced effect.

In 'th c1w1:.re cx_~cri:-:nts, 113i*¢,a:ii:\xren+ 1n1'“i
(l 59o) C\ul4 denonetr-te rc netrrxent-l effect c
e“*lgin eiceee r’tro en.

ACCWrci’g t? Jeri-v-53» Cm (1757), in rev

SCiIS. 339d
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re grown on neev; rumic evil: re her
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- ‘ ‘ (1 . 1., flap , i V t i) -,_ 1,. _' " ’.,. _, ,1 l
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n n rnL4an we L'n w.“ es+wd ”a e
1;;IaLlir'1‘-_L‘-'1- (FL LIN; .lctl V531,: seem
er ccwu rt ens (tir1n true flevclf)1t3ni

ardei as very

‘
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beet seed (R APP 1953). b;

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ede urict develofed and natured a

Q .3

la ~z

ure were sifnificantlf were rapid in
90 C mean emperature.

’fttle is kno r abwut the e fec: o?
it“ or er inatton in en ar meets u

did not

 

archers (Aiflth 8+ 31. 1957) re‘erte
in: less thfir :0 her ent 0? Meter nu
4 train; 3 ilfiefliat3l(. SeofiwaLIT
Cevflieff. ’f‘ZTV’ Ed ‘i?i:lr Vlsll 9“ in
in ar.3~ o? stire e. Accor”1r to
t cl:SYCi"vaces (:‘olfi 1»: it ,e3‘=a’ in
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capacity drOpjed considerably (10 percent reduction) when seednalls
contained 75 percent water when harvested.

In some cases insect damage, due to different Lvrus species,
was ooserved to be responsible for a reduction of germination capacity
(H1115, l9h3). This occurred during the early'steges of seed develoo-
ment (HILIS and TAYLCd 1950),hut could be controlled effectively by the
aoplicaiion of S nercent DLT (HILLS et a1. 1916).

Apparently sugar beet seeds maintain their ;erminetion cagacity
for a lonr time when stored under dry and cool conditions. Jeedballs
stored for 22 years around 00 F still germinated 75 percent, only about
10 percent less than the original material. The resultin; seedlin;s
showed normal vi or and the roots had normal sucrose content at harvest
(PACK and CHER 1954). At room temnerature tne iermination capacity
dronned rapidly after 15 years of storage (after 10 years 96 percent,
after 15 years 59 percent, after 19 years 27 percent germinated). On
the other hand, FILUTOVICZ and RRJnAd (1954) observed that seedlirgs
from seeds older than four years emerged 1-2 days later than plants
from younger seeds. Under storage conditions of high moisture and rela—
tively low temperature the viability of the sugar beet seed is quickly
diminished (PeTo 1950, HUhTER 1951).

Different seed processing methods (shearing, seimentin ,

decortication), used to decrease mechanically the number of true seeds

er seedball, can also have an influence on the germination character-

".1

istics of the sugar beet seed. TOLDAN and STOUT (lth) observed that

V

sheared seedballs have a lower field germination capacity then tne

rcr al ones, because .7-15 H.;rce1t of the smeared seed verminated
abnormally dae to the e"Ԥs ire of the true s; ds to air. hlole seed-
halls and se vented seedoalls yrocessed to the sire size {ave HUJUL the
sane germination percent and e er ence results under co naraole field
conditions (:13? lQLQ, BLufifilgcn et al. 19h9). Decorticated and non-
decorticated seedoails were re orted (infi lQSC) to have essentially the
same field germination cuoacity, altnou h Lgacn et .(19 a: ) found t: at
decorticate d suvar oeet seedcalls :erminated and e erbed faster at low
soil moi s+ ures. This benedicial effect was attriuuted to the increased
water absorption of decsrtiCated seedballs. at hi_n soil meisture,
only 51 i ht differences could be demonstrated.

Polypliid sugar beet v rieties produced artificially by
colchicine treatment possess generally a lower germ nat ion capacity

and lowe

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the correegonding diploid ones (iJTC
and HILL 191e, Iv’AI‘jUZ-ljna 195:1, 3n)‘.)I..I-..~1"1’;t 1957, R3}: :1‘.Y 1957, amt-TL et al.
$1957).etran13id segdhalls, althou;n larger than diploid seedoails,
coatain fewer viacle seeds. FgTSTv ; (1,5; su;;e;ted that the

inferior gerhihation of ooljploios is due to meiotic irre_ularities.

£.nTL et al. (10;?) cauld not confirm this finding; tug, sa__ested

. u\ '- ‘ r' J' ‘ ‘ - 1 l 4' . ‘ r . V ‘~‘~ '~ ‘7 —~. "‘ " I ' ‘ .‘ ' " 1
instead lflnb the re acivel; lower Lu oer 0: seed Lhandefo 4L1 the

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Jan 91" n11::.oer OI. 1311:"...3' 8377'} Ci‘ch_L:-.; .Ll’ e I 99‘1J113-U.._° l 31." C 9 11”“: L‘: I" c I"

germination of polyplOLds.
For several dec - fies, intensive investigation? h~ve oeen wave on

the in: HlU*nce oi' seedhail size on aeraihativn. ,nie great interest tas

\ g

due t3 an early re art of oxlgr (lde, cited by AiCHl:;VIIUn 1950), who

claimed that larrer seedballs pro uced not only better field *.rwinatjon

(J)

reauizs, nut i“ve rise ti were v1;wr us y1an¢5 2n‘ hi_ner 31eld *Pzn

‘ 1, w. L. ‘1' ,- , 1 1 - w . ' ;.' , n " ,» -

tre Shuller needr3110. ”Lihsu n =11; 3 Flh-lr S CAJLJ rct we con-

“. ' ~. _‘ n 3- ‘ ‘4 "I r'\--’: "~ vaV .— .\ r‘h ~v-lr. Aw; y- L. ' (x, :1 \v‘ r r- ‘r r A (“r-w \
111‘ {rid (QC/Hr. 51-1.5.3. 17,12) 11 H arm: '1“--.- u : .I‘-. an. lI/HH’) C11.L1‘.‘.- 1.) \f F‘ ’11 o 151/ I/ l )

a positive ccrre

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thls corre1~110n 13 Lne uaxnly t0 the peorer Ferminatinn of the e a11egt
I 1 o m ' - u' (A v ‘ ‘ _ ._ . ‘
Ceedball c1a:3es \;1ameter L¢jl1e Erin 3 mL). Jer; large 3e91u;313

have no a vent 3e over Kindle-eizei rte: (ILCL;L" 13.9, SEV11441 and
CCX“L?C$ 1038, FLICfl and CALJLJL L§Lfi).

In mu1 iterm I'ruits the problem of seeflball $17e i? very mLch
COMTLTCQLGd cg the fact thct 13r;er :eedsall: c:it;in ;r3;“rc;uwtl1y
more true seefi: Trev Era;Ler (neS, and that Che size 0* tne true seeds
within the 89:0 seedrall 13 11:0 verienle (0:1LCJLLY et 91. 1757).

‘1
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(Vrfi' '\ r‘v\ _‘_, . “‘ r. - .. ‘fi ~-.
Accarrjing to JLANL (175;), true SQCQ; have w;y'2(Lma1u1» the same

Winq+ien cayacifiy whether they ceme frem 3QG, its, or xu1fieseded seed—

ba11s. However he observed, that true Seeds 1n the sage eeednagl de rct
penninate finde' .nme _t1y, out tnrt there axiste a certain ”cmnxunjfv of
fate" between their germinatjon i.e., when SeefinaETS containing tv: true

seeLs germinaze, tnea wreuuce eltner 4 seedlings or none in greater yr)-

(n
4.
n
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portion wnu1d be exrecfied theore*mc1*. Th13 hex true for inhere-

tory as 1911 as for field cyrhitions. Acevrdin; to C LC€JKY e? a1. (1957

 

larger true seeds produced seedLinga with better vi 3? Ann heavier rnots
len did soa11 s eecs from the same seedual1.
" I 7?-' .I'.‘ ' r 1 \ 1, 1 f‘. '4. .‘ k 1 ‘ .-: V‘ r h , ‘ "1 ‘.. ,-
DHVLPJHY g} e1. (198A, TaJLe 4) need true urea. ”L «Onc_crm FLLLr

V 0 v_ ‘, . ‘ ' "I a'. ." . \ _ ' ‘, 'A‘ ’ .'. .
meet 1:ne3 to sunny tn. e Lee: 01 frdlt ‘128 on yrwlrufrnh. LsL1e he

sifnificant dinerencee were dehonctrated JPLVCQJ ;erm1n Lien of fiLe

eif?erent none-err frrjt c1er<e= a.e11er Prmit: :QLeteu tm 17T51nkf€
Q

somewhat errrLLer trig) 12r“e area. a'aaL'y recmremui that, Luvfier F5e

conditinne lerre menoyerx fruits gzve a tniwrer ET the: Le4Erx cr

snail sized Ones, e7*nfinfi" the O”§O§ife cannLus jar was drewu in pre

meets. HST? Afob (195'?) ”“”01 tr eted c1eurlg the: Levier monAQerm
fruius bosee'sed a 1ewer germination capacity and e elnher rete ef
emergence than Sma11er FruLtS. gunujnf of? same of tjg pcri'arp
meteriel icpreved g rwinatifln cnr:;aer~uL/. He convtuc-d, therefere,

4‘» ' 1“". w m“ r -"~.w .0
LURE the _re;cer dg'upt o”

VQ“1CL?W m912r121 niecrzased «urxinatimn

.. —. 1 3 1 . ‘ .74. N. ~-\ " .vx _’- ‘. "i I ‘\ \‘ f- - ‘ , ,. v"
c pecLEI and seed11n, ngur. me Ju~993teu cevechlng Leno erw eu_1r

-. L18 531109

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as a Keen” a” imprevinf ge'anniwnr hgrkjnv Lian wnnngerue, 5:51 wa
flan 'V- / ,_ -r ‘ ' .9 -. V. , ,1 ‘ r. ..
and :Lfl' mi. (1900) found aluo that, excewt far tne verg r LLle:t aiZGd

fruits, sceed of yenninétion we: inver5e1y correth 1 En the 5128 of

1.

fruit (including bohn diageter and chicknees). The Fruit* of medfium
Classes COL 'Wned the hlghesh progcrtinn of viah1e Seeds.
The re1ation3nln Detwee: eeedbell siz, erd rerujnfition is
further cnmpLiceted hv innjbitory subetwncee eresert in the fruit.
According to DUYK, et e1. (19A7), JCATYS 3n 1922 prcyared an extract

from en ar beet eewmthLis and recoirized its 11H110iLOPY eTfect 0L tne

true seeds. he ale: reve the lirst are1yeee of tie inor_zLLc cw~poner*

of ”he e extract 5. LU: c,1n (1939) dewnnstre te d the, the Water :O1n019

inhibit0r(g) are “J9 SF€Ciiic t0 the 3“eciee, but returned Lflb ermirL
tion of at least 29 other Species as w911.

Independent of J3 TVS and FHCLBCEHL, TCLALL and iTCLT (194:)

prnved corcL uvivelf That the corky utterial of the su;ar beet seednall
(periCirr and parianth) contains water soluble suestances ret-rding
gerrinetion and sometimes even killin; the raiicles of the seedlio s

in laboratory tests. Extract from the seedballs of slovlg Lerninatinq
seed— lots retarded tHB genuinotion of the true sugar beet sccds in
comparison with true seeds 5 rminated in water. The great differences

in the percent germination of seedoells of v rious varieties or seed-lots
almost comolete 1y disafreared when true seeds were used as a basis of
comparison. They concluded that the inhibitor; substances, waicn were
present in the seedball, must have been responsible for the differences

nation, at least in laboratory tests.

H-

in speed and total senr
The* suggested also that the water soluble Lermination inhibitors
vary not only with varieties, but are influenced by climate, soil and
seedball maturity as well. TULEAA and STuU1 (l9tC) assm ed that the
toxic substances interfere especially with the forecast of potential
field germination as measured by a laboratory test, because unoer field
conditions the ab orptive newer of the soil renoves these suostsrces.
On this basis t1Lev recom.Ler ded the washing of seedoal1s in running vater
before testing them on a blotter for rerninau on (3-,t1 and TUL An l9ulb).
WLis nrocedure also decre sed the rercentape of "browned seedlings”
(injured radicles ?) ( AlbinShw 1958). Accoro Ln‘ to to 01f ial seed
testing re Loos o: the United States (T 911“: AUULLLL LAAL ALD JsLU-A 13
SEEDS, 1952), sugar beet seedballs should be soaKed for 2 hours and
washed in running VLater to remove Mater soluble iLuLibitors, WHlCH may
interfere with the laboratory germination test on blotter.

Very confusing results have been rWu blished in the literature on

ll

the nature of the presumed inhibitory substances. STQbT and TCLLAN (lQLla)
claimed that in dry seedballs the germination is inhibited largely by the
toxic action of free almonia, which is liberated from nitroaenous sub-
stances present in the secdball by enzymatic hydrolysis. however, in

fresh seeds they suggested osmotic pressure to account for germination
inhibition. RENE (1953), using garden beet seed, also claimed that

free ammonia acts as an inhibitor during germination; but he found that

L. ' F‘V
l

the ammonia was formed mainly through the action of bacteria. herefore,
he concluded that this type of germination inhioition takes place only
in laboratory dishes, out not in the field and may be prevented simply
by the use of seed disinfectants.

DUVF, et a1. (1947) reported that 05 otic pressure, due to the
presence of inorganic substances in the seedball, retarded germination,
but they were unable to obtain any srecific substance, whicn inhibited
germination.

On the other hand, as early as 1940, GILLIj fzund a strikingly

high concentration of oxalate ions in the beet seedbail extracts. He

dewonstrated an inhibitory effect of an 0.03 normal ootassium oxalate

(+

solution on germination (cited by QUYM et al. 19A7). Independen of
GILLIS, NALIKO and NIYAR‘TO (lQSL) found that the most remarkable in-
hibitory substance in the sugar beet seednall was water soluble oxalate.
This was demonstrated by KIYAKOTQ (1957) for both dry and fresn seed—
balls. He claimed that only 50 mg percent oxalate is toxic for the
germination of the sugar beet seed, While he found that the seedball of

the investigated diploid and rolyhloid varieties Cirtain about 2 percent

oxalate (50 times higher than the toxic dose). Seedbails of diploid and

F-J
')

“ol' 1"? v riotie: ii’ ~at di ier 3; if 101Vilf in i11:;c ac- t 4 wt.
F? KC73,.::”:1. (lofl, 795;) niinihsd fro; the water criract of

the beet seedtell an ursaiirated yellow oil, w ich acted as a titerjol

garninition inhioitor when tastei on doth QI‘BT beet and other Seed .

filth u‘h TRCY THEE (1955) adniited that the W,rrinaticn inhioition in

--~‘J‘A‘v

‘ l

lets i laryelv osmotic in nature, neveranelets he c uld aeronatraie

.-. I-vi

that the demineralizated extracts of the seeioalls xl‘o innihit jsr;i—

natiOA. He conclude” that av

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tnere‘ore, rust ce 8-tFlbu.
A3~r +‘ ’33 ~r\n f”. ‘p‘fiI‘VW "‘s nr: Q-‘l ’3 + "- V"“""‘+(\" ‘" ' ‘r . L) I ‘ "\r‘\l“(‘

.‘g‘ Jnxt tic)"; 0.. r er... 8.. \'—L\-.~C\lr‘VQC.AA 'v ‘1‘AHV‘, I‘Vi 111.c‘A L ..'e~.-!-l‘- ND

lfl%7) fwund that o~01foenzoic, vanillic, p—orgcinnawic

an. fernlic acids in su er beet seafltall extracts inhibit ternination.
He also isolated ferulic and vanillic acid from the fruits of the other

isceie. hecently, similar results have seen re dried

 

O.
by Hungarian investiiators (KQVTS and VAhfiA 193%), who found p—oyyhenzo c,
ferwlic, r—couriric, caffeic and salicylic acids in the pericarrs of

T‘~ “r
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:ev also recovered thes comncunds from nunersus

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other fresh fruits, as well.

On the otter hand, SLYDQR (lQEOa) presented eviden;e that, in

some cases, sufar beet seedhalls may stimulate :QFMlnfltlfifi rd early
seedlinr Frowth.
Alttwunfii it is genierallyr:assuteul that ixfiiihitcnjr sun t1 more are

innortant in nreventing a prenature :ernination of the seed, nwtning

1...

definit ve is known concerning the infortance cr role of the ore ugso
are in the sugar beet seedhail. accor in; tL DA hcfl ,

et al. (1953) the Presence of a water soluble derrination inhibitor in the

the sail has attained r sufficient {disture ontent, b; w icn tine the

ttef ‘teculated

, ..‘ u ‘ ‘ ‘ ' V. t r‘ ._ I: I ' ' I ’ o ‘1- 9 ,~ g ‘ ‘- v 7‘ . " l‘ a.“ .r‘ "I
suséest tilt the nreeeoce OJ germination lflhllloOPS in fruits 1: a ;e»erii

rhenonenon, which may have an e sential role in the biolo v 01 the riant.
The nresuxed impartance of Lernin tion inhibitors has over-

shadowed sosewhat the long recognized fact trxt the bhgfiic l FFOjfirtiGS

O

‘- r, A. ,: ., “.1.., ‘1,_‘._.l, 7", ,_ ‘ ,w. a: ' 4.‘ . ' ,3- - ‘.
f the see; all it at: gay hinder Vernititton in the cultivated so er

(I)

‘ 'L' x -I " ' ." . 1' - . ‘V 1 I L". - a ’1 ' .~ .« r- v r
beets. The totrn vood-r nericzr» ard the ti htness o: the seedC‘y Mare

0

sur ested as the cause of non-vermination in certain Beta subspecies

(5TELRIT 1950, 35 SA 3957). From this it is logical to susfiect ti t in
. .. . . . . . -. -. ~. ‘ .r... A. / praw _.’ ' H r ,
Vets vul_°ris similrr difficulties nay eklbt. ag’Ufim leDJ) in a search

 

for factors resoonsible for the delayed gernjnation of su or beet seeds,

observed marted difference" Leiween and within varieties id the tnicx—

(H
'3

ness of the dense inner layers of the seedballs, as well as in the
1'

the seedcaos. He also demonstrated that notched stedoalls

o -o 1 \e . ‘ .: ‘ \
have a faste gernination than the non—notched seedoails (3; out levee).

f‘
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On the other hand, BFATL et al. (1957) found no differeyces
structure of the seedball or in the ti_h:ness of the seedcags between
the seed of colyploid and difloid sugar beet VSPlGQlGS.

The fonerally recs nized beneficial eifect of concentr«tej
acid treatwents on the :ermination of sugar beet seeds can he rexarded

also as an indirect evidence for the nhysiral hindrance oi the seedoall.

Germination could be significantly improved my concenerIted H 30A treat-
went , which corroded away much of the herd pericarp material and increase
its pe‘aeability (HiWLdY and hCQDRAK 1930, ANOXYM 1931, GAMLER and SA IDEIJ
1932). Concentrated acid treatments were snown to h.ve also a great

Ffect on the loos enin of the seedcaps, which are cenented to the seed-
balls by heui—cellulose (LACKEY 1€h3). Due to the above eifec ts, tne
concentrated acid treatments increased germination capacity, hastened
emereence, and produced a greater plant population at harvest, which some-
times increased su ar yield (FALLADX and hm.1ounuH ER 1916, GRACANIN 1928,
HENLEY and UOGDIAN 1930, fTO'Vé 1931, GARTER and SAKDEAS 1932, dARTL et a1.
1957). However, according to TGLEAN and STOUT (lQuC), the beneficial
effect of concentrated acid treatments may also be explained by the
removal of an inhibitory suost nce from the seedhalls.

Besides concentrated acid treatments, which are somewhat dangerous
to use in the practice, a great variety 0? seed tr eatuents have been pro—
posed in an attempt to stimulate sugar beet seed germination. EIYAYQTC
(1957) found that soaking the sugar beet seeohalis in water solutions of
different inorganic and organic salts increased sirnificantly the germi-
nation caoacitv and sneed of eernio tion in soilt ests in the laooratory.
A 1:1 mixture of N/lOC 0 III(CH3CCO)2 and O. 5 nercent NEJOL gave the best
results, especially in the case of a tetranloid sugar beet V'rie*.“r
NIYPJ 0T0 (195W assun.ed tiat this eifect was due to the inactivation of

/ I
the oxalic acid content of the seedcails. FANUY and PAL (1958) reported
that the annlication of 0.25-1.00 percent KNOB solution improved consid-

erably the seirrIination caracity ofr nolyoloid sugar beet seed. Sut

PJ.

generally it was found that soaking su~ar o_e t as efioi is in mater alone

15

laaltwmiLn ‘ he I °Tl"l‘tlmlh (1- tI( KL 1r riaet 2338" ('C1. a. arv1 J-‘I “ l‘ 1U,
n n1 1 v '1' '1‘, 1“: - , . ‘ '_ -, ’ ,‘ .'
olifI-l- 1947; i. lax leglj. I vever, 1n f1:ld e- e.1 en 8 no :l‘Tlf-

c le oe demonstrated vhcn ;hc

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8419:1921 L.) 1fi;er:'3 1"“: l1.[ . 61 0 r‘l’”. 01.333111” I‘gil‘t (39'3“ 0,5; L .13 71197‘3- :1.§f\1 tr .J 1" .1, C {31,
I -1 “ C: ‘ . ‘-" _ _j -~.——-~-,-U .

. 2-1. is? _21, Ht; L3,}: an.) 1.3.... . 1L 5])
- r‘J ' ,. 1- r a -. ' ' .‘_ ‘ \0 l 0" .4; 1 ‘ -, ‘ \h _. _ ‘1 -‘. '.: ..' ,.
r€.1gln.. 11113 3882105111 .3- l’».]_;,_1 QlIffl‘SF‘VVlL, “011,0“??? “dd T10 [19731791. 101:1].

effect on gernination, seedlinc ener: once, or seedping vi or tflgnjyi

seedoalls siwilarlf had no ef'fect on ;ermination, vi or or yield in su~ar

= “of ~... 2 (-1" r 4'.) I ‘7‘” ("I " \ 1 1.": *‘ - -’ "g".-

oeets (U .I’. .~\ [Lit 15’ ,II ', 1.5-1:.-JKIJ 1 75.), 3 . 1’ 3:7, 1‘:-’_‘«;‘d.j .
a ' .- 4‘.» a c - Tn ‘ a ' I. ,7: ,...-'
twatinr u.e sur ace of eukar eeet :‘GOHQ ls 11th hytrQLH11-c

col oids (felatin, a ar, and "al in") accelerated u-ater unIake 133m
sand and accelerated QABTfBfiCS rnder fiell corditions (EIVJTCTQ 1959,
DEXTEi and PIYA.-ZC 1959, NIYAFLCG and DEATEfi 1920).

Relatively few attempt :een made t1 infrcve the erxi-

U)
:7
As
<‘,
ID

V

’ '1’ ' ' ‘ "’ . " V V ~ ' ' r. r J 1" ='", °—‘ 'm
net on of toe SLFwT oeet seed 0; _1ant breeiin: methods. Luis 10 not

for assunin;

U1

too surpris in since almos t no defi Ini tive eVidence exist:
that L-rninnfion characters in sugar beets are LGhetjCEllj controuied.

H308, et al. (1950) and 5200 (1952) have shown that uncer low

0

J

temperature conditions (2-3 G) varieties differ with rescect c1 tLe
aOility of their seed to germinate . hr Srwnle wass seleCCion Lerninaiion
vas improved 20-30 percent, showing that sufficient ;>re ic v ri: hilit"

existed between and within lines of sugar beets to reryit selection for

this characteristic. Diiiererces he seen varie.ies and inored lines,

1M

‘ . " I 4' ' r a ~1' ' ’3’ '- , i»w'? - ‘- . '~ V r» "2 ~- ; . z~ Ixr ‘1 w , :1 n ., ' : ,'< \ . J - - -'«
M’n1021 113:? V“") F Te EC) J ‘ 1. " 3.;01Lr' ‘1'}, 1 ! anA .2 "4&1 0,1. ‘ ”.1‘.,.,L,I.L.‘ 1t-‘_1 .1 L" “”3

~" :r ‘L - ’«-~ ~ .--~ , ‘ , vv « r’ I ' ~ r; , , , u. ~ .
+euireldzxgre, 1 :re .mrtz 1“Ler;'3n9» Luere 1aeét r51“ '1 4— , (J t::3«3r3txx1e.

Oh the basis a. selection e_jerixent3 311; (LQ5") Cfirc need the; Lhner

(-r

low L‘u“~ru4rr" Cfifld; iens zuotzi rate 01

lihgs seem te be h riteele tr its.

 

‘ 1' q Tm a- i ,: ‘
diploie VJF1€;leb cu- 3e votei ea.

rvw "5‘ . 31 '1 r hm.‘)\r1'— ,
5g\bmx ohse rve ed lerf eiiier:aces 13 race of \ffe1h3uitn wt

D
:l“
tn

of different varieties (1935) and in ener‘euce 01 such seedlir s
from $331 (lfifih) or 5011 (I? 7). There was greet ci'Ference betveeh
seeflba1ie harvested free irdivifiu 1 r1 wt: of the Same variet: vben
c:rvin3tefi in certect Vith nutrient solution of 10.1 3M 'hzres

osmotic prexsure (SAYLLi 1Q97). he has SU{,CTted that the meior i We“ r:

I +T~q -
arm 1 -LDJLUS

O
O
3
(4"
d
1.1
._J
3
L;
.J
m
(T
ad
0
‘W
I
H
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a
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0
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D
(‘f‘
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(11'

3. 3’1 (lQKOb) selected ragid eni 310w gerniha+ih> seed1in-

from the see 3 of a br03d— e3e, oreh-poilinated variety. r1 nts Hfltvin

the two greens were allevee t5 inter-Dollinate and seeieills frei Y2 of
the re id and 72 of the slow Lefiminator e1 ants ere ceq_are” for e~eed
of germination urinj a livsid.— —c0ht act .srxinat on techuipue. tn Lée
averlfe; +5“ 8' of the r‘riw ‘wrk ne+'h. lento anc‘ed “5rsxdct

f3 er enmivation th:n t1~“e fre th5 C 3% w“ 1h“ LL .. h, . k‘ fire

1‘ . r‘ - . _~o—-. I A “V“ j . V 't‘ 7. .E- »~ . .~: ,. z r v» x. .7 x - I‘ I m.“ “ ‘ ~ -‘
Helif an 3“0;LUV1 ;ciuf10n, mHlCH uIOuthJ a wut JO qeLL;.hepeQ

osmotic flépchfp wimla.gm ani flhh Jl (195d) bade :818CCJUM; dim re—

,, a- -
selecticns for early edrwufihy and late Sprouiiné seed inL: in iwur
comrerciel sugar beet varieties. The earl; s renting reeelectivns

from all the feur varieties germinated significaLtly faster then fine
late ones, whew teeted in inogiiol solution. Due to tne improved syeed
of germination, the yield in one out of two yeere increased signiVi~
Gently in comwerieon to the ori*inel parental varieties (QQKTATbn, et e;
195%).

To avoid refietition, literature cite ions pertaining to the

applied tec;niques are given in tne abpropriate sections.

 

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Vite? 1N CICSGJ Fl::J.C 00 9:. [n eicw LJA a Wiece O; fidrdh screen Avg
.V V._‘\‘\ (5‘: v o “‘ I1 ." .’ ’. l _ ‘n . . r—l V. N :» ' “ v ... -
eu'nwrted on live ‘lJCQ> Oi ~'i S cumin . CJQNC' SEQCDdltw vere 4:«00d

iii eqc?2 cvaj“ler' f time bCX’jJ1 cetfiwqct vxifih iflie xiirfacezcif 3.9“rcyiufzpeflv

Q ‘ f1: ‘pflfi \ -: -, ‘ -.r -VN : w. x-» ‘. .'.. u: .. ‘ F1“ ,
laO Kiliiiite? of a b ,erce- rirenal LqunfimL eeiuusun (L"uT€ l.}. ive
Miler Cemfimsiiier a? the nririefif :wlutinh vac a; fellfiwfi:

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n “7' I P E o nr\ 1'“ 1' I“ - I . 7‘“ _ vA‘ ' ;’ '- . -_ i '» ,“ _r‘;4 '.
C31C» .9. ‘4”.2“. , “~-’0 \- (‘13 .1 '9 U‘l~(\."~)01{+r wk” C). -— «14'1" \‘JJ. , O. .IAL‘Lf) , AWZF‘JA, it. [3.1' ’,

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3 )C .7u’jk." ngltejpg,‘ :‘.-i..;\i.,'bo_.../,"/‘U, 0.x»;- Do if“? C‘.1r;.'\."u] C greibul :; C! . 1.. .3
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solution 3b lb.l afuesflereu. .0 aszJ Vdrlaulflh 1h 39Tmlflmt.ifl M A~.n

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dried in a plant drier at ,9-j3‘ C, and efirefined my a anquagjpn

threeker ind drqrev deti"e t d t“ wehflle $331} :e.gie3 («{U;:Zzgij» **d
31é1a*3 lQLJ). 2ffe” threthin; fifie ace ball: were simwel afi viva
conriii WM: for at 1‘? fl;&3‘zee“ me 7 'é*,~§"uere *e Tra"“v" 0“ :,,r; “.
l'fi‘-fY‘"(‘P’ ~0- r35“(1’-“. "'v-"i-fi u r i 'x :1 1 ‘1'"3 3‘“ ir" “ f .
The xéeflfiélls were re: freefed ViTw a fifivicifie. Q&-;.;u‘ >L“ 51:[
(7°53) reveried *Paf lrr'e” éee*w'ile We“e leis a‘fefl;ei wf ”‘ ‘7“
S"1l‘.111~r‘c i. } ;"‘(3,q ~3‘,;~ ",-).n",.-‘ Wfi'l '41‘ CV" ( l 1"

 

 

 

 

Fifiure l.

Top View of plastic box ucel in liquid cont
technique, b
and {lass rm
tch of the

 

“hcwiui arruh’ewent on seedrqlls
7"‘V,

.I.” r-id3‘

 

ds snvtariirL the screeN. tunir 9 on
screen

bcx. Etch seedlfit in a geniimeiicn box is r»
seedhalls.

S€“9*Ete$ Tfiur d;ffereht eeedlnts vitrih a

 

“resented my 20

/ r -. / A g v, ~ r‘ .x ‘ .' ‘ ‘ ~ , ‘ V. r‘ ‘3 .‘ ~ _ c A 4_ n k _
ll/Oh - lg/Cu incn (4.@ - 3.1 Rx) die eter vere Q;UJ. 1 ,ii WdTMBHQLth

\

rercent was also determined far e30:

(K)

dldt, 5n that those with neor

U]
(I

viability (less than 85 percent ieruination capacity) could be enciudee

from the results.

-" . :l. 'V ‘._ -' ‘7. fl,, - r... l ,.~ V .V'- - ,,
All {QVAaLdthfl ehperlwdflts, excegt d; rdted, “ewe cauuucied at

(D

b-‘

room ternereture (22-250 C}. L “wingtion ccunts w.re nade L?, 72, and

1:0 heurs after the 3 ede were rlaced in contact with fhc nutrient

(.

y

L
‘
w
J

solution. A seedball was ceneidereu a; {iiiinated wreu inc redicle of

~

' ‘ rm

at leaet one true ewed entended D gnnd its Seeccoat. inis could be

decided onLy by a cere?ul examination of encn seednail, and Oftcz only

Genetic-statistical retho‘s:

 

J(H (lCCFO (Vanniatrated +1ul,()47' envirarwfadtal and
genetic factors may contribute to the ooeerved phenotypic variation
between the individuals of a sefrefitin; population. Vii 5L ilvla)

1.

it useful to empress this relationship in the folloving linear Form:

em; —4
H

2 2 a 1 v 0 0
+ ¢+ 6;; where 6; renre se _ 15 the total observed pnenmygéic variance,
q‘ {:3

.J

65
GE'is the r.art of he nhenitv'ic variance due to genetic effects, chis
the part of the phenotypic variance due to environnental factors, and
is the variance due to genotyre-environmental interactions. Elddgh
(1919) further partitiored the [enetic variance into three cougoncn.s:
(1) an additive genetic part, associated witn tie avera¢e effects of
genes (63), (2) a dominance pert, arising from the interactions of

U

lleles (65): (3) and an eoist4tic part, a triautaole to the interactions

m

of non-alleles (0%). He also snowed how adiitive sene+ic and dominzurce
variance contrioute to correlations among reiet1Ves in randomly breeding
ponulations.

NAihiyi (191S7l poi ited cwfl;‘tne diiierfz ce C£3W¥i ci;;fixexiie aimiiin—
fixahle genetic variance. Untixable adl'iive Cenetic v riance arises
due to the presence of dominarce when the allelic gene fT€(H€HClBS are
not equal to one-half. Recently, epistatic v riance has been further

cartitioned (C ch,;:nfl l95t, Kdirlrozig 195L) into factorial con;cnents
(additive X additive, additive X dominance, dOMlilnCG x dcuinance, etc.
interactions) to estimate its inrortance compared to adritive :eretic

and dominance variance.

p,

(a) J

LLSH and PthE (lQhU) prohvsed the ratio of the additive genetic
'1‘ A
component 01 v rianfie (6:)'to the total onenitpnic variance (Uéfi’ a: a

measure of the degree of herit hility. Later, LLS’ (1943) referred to
this as to "heritability in the narrow sense"; whereas for the r tio
n , 1. ' ' . 9 .— ‘A + 1
01 the tOtal genetic v riance (6? - + Gé'+ -) to the total rneno-
(17 D n .L "
tynic variance (5%) he used the term "heritabilitj in the broad sense”.
Heritability values are exyreesed usuallv in percent. If the

U

estimated heritaoilitx value (hd) is high (close to 100 percent), the

‘1
LI

observed variability must be lariely due to {enetic factors. If the

C)

heritaoilitg value is ion, the observed v riation is nainly due t
environrental factors, and little gain can be exgected for the given
‘1
.1

characteristic from se ection in the particular ropulat on.

In the way heritability is defined, it is implied that — strictly
speaking - any h2 estimate of a certain characteristic is valid on y for
a particular population and for a efecific environ ent. Also, at present
no satisfactory method is availaole to attach confidence intervals for
heritabilit; estimates (KEL3ThCin; 1957, thsoN 1959 . Despite these
shortcominrs, h ritability estimates can give the nlant breeder very
valuable information to predict the gain From selection, and to deter—

mine the most effective method of improvement in quantitative charac-
teristics. This is estecially true where, for the same trait, Lore
than one h2 value can be estimated using different populations and
different statistical techniques.

Several different metnrds have been used to calculate herita-
bility values for a certain characterifitic. These hethods vary accordin

E -
n.)

to the aoolied statirtical tecnniones (re ressions, covariances, variance

components, etc. ) and accoriing ti +he nature of the pofulation, from
which the data were used to estimate r‘ritaoiti (inbred line._, ranr>11v
breeding nonulations, etc.). hitn the excention of one method (estimating
h2 on the basis of a selection ext riment), all estimation methods are
based on reL 10n31113 anwon relatives (parents contrasted hltfl off-
springs, half-sibs, Fz's, Fq's, etc.). No attempt is nade in this thesis
.1
to review all the existing methods of ootainin; heritabil it estimates.
The sugar beet is a cross— oollin tin5; crop with natural barriers
to self-fertilization and controlled crossing. Therefore, the classical
rethod of esti misting variance components (hAIhEn l9n9) could not be
anplied directly. instead of this it was assuaed that a commercial sufar
beet variety is a randomly interbreeding (nannictic) ponulat on.
Genetic variances ano heritability ratios were estihsted (Exyerinent 3)
from such a nonulation usinr o .rent-off: irin} rerressions and between
and within (maternal) half—sib rel tionSLins. The oarent— oiisyrin;
regressions and the between half-sib family variances can contain only
additive genetic variarces (6%) and some epistatic variance (6&9
(C0313 nal 1956, K‘E PTHQRJW 195? ; the within half ~sib family Viciarcrs
may contain a certain amount 0V dominance vzrizir e(6:) as well (ihfiH
lQAQ, l9h9, LA hSR 1 A9). Folloving FREY and Hofihfli (1957), instead of
rarer t—ofi‘sjring5 reeressions narerut— off:*r1n correlation coeffi cien ots
were used to estinate heritabilitV.
As a Cii‘Ferent nethod to estimate heritability r"or speed of
re"a1retlon, the variance between clonal glants was used to e tin te

the envircnnertal variance (Exneriwent C). Co marin” the v ri:nce 1)-
I

taired betWeer these reneticallf unitirm jifints Mitn the total variance

I \5)
\D

h2 valves will C”rresnond t1 h ritaoi it" in the broad sense.

' u

As avotrer method, h“ ratios were estimated on the basis 0? a

n
1 I \rwxr . , " 'lfi‘f .» ~- T“ \. z- .-. . . 4 ' (V-r -' - ' 1 \v‘
se:ection eijcrimeno (nmnerlnwnt J,. Tfi Cwlculite h , the 5a1n {11m
. u u 1 1 J51_ 5‘ q .5 1 ‘ 5: “- n 5 5- -5 n v ,' "1, ,~‘ :5-
selectton W33 11vx em or the s lect or d1 ierential (L h.;fl 19,3). in
trig C359: :eLec ion H3: fiade in the th1 Os osiie direct OHS, thus rakinr

the cornerison oetveefi 1%: wear: 1? the two selected groups and the

c 5- r5 5 ‘5 1 rx 1 _: 5 - '_‘ ‘l 5..., u ‘ ’5 ’- V - ‘ . 5. z“ 5 5. a 5'? r v I 5" A
or] lflfil rotiik ion trwx;ce::31fir. To 61.21 J? btfflfl):nlllcd= b5 2 1.
:xethod, the as olute iitference between tiuzzwaan: of the t‘: selCCted
‘ «' ~ ' “ fl} -: i ' I . \ r~ “I . ' I ‘ 1‘ fr "\ V" : ’. . " ‘r'x "' J 1' " L" , ‘ ~" 1
£3101.th oft/1r 4rterrmllijh—H, {(7,3) (nolycr-ifla) 12-1 , 1" 1V109-i cu, v.19 _ Logiiut-e

: in M, ‘ $- 1 u ,5 . 4.‘ ', ',.‘ ... - '. . 1 ... .1 . ,7 .2 ..
H"lerw~ce be weel the Le1o: of ere ori 11 1 th. selecieo no ilatgone.

.r...-A. «A; 4 I v’ . - V ., -- . _ . I, .5 . . 7 5
, A A

EKFfldlzyffnL daiULTS

:1 .t-‘
vaervment A

_A

To determine whether the levee differences observed in efeed
of germination between seedlots* are due to fQCLOTS re idin! in the
waternal tissues, or to factors present in the true seed itself, the
germination of seeds in seedballs and true seeds of
were compared using the lihuii-contact germination technique.

The seedlots (harvested in 1957) used in this exnerinent cane
frim individually harvested plants of U5 401, a commercial, multijerm
sugar beet variety. The seedlots were previously evaluated for speed

Of germination by Snyder. Out of this material, tho ertregely slow

(118 and 12%) and two rapid (528 and SA?) germinatin5 seedlot

U)

were
chosen for the experiment. From each of the four seedlots, 8O true
seeds with the brown outer seedcoat (tests) and 80 true seeds without
the brown outer seedcoat were obtained. (The true seeds were renoved
from the seedoalls with the aid of a fin5ernail clipper.)

The exterime t was arranged as a solit~snlit plot experireni,

the main plots being the four gernination counts (made after 2A, LR,

 

*The tenn "seedlot" Mill refer to seeds narve tea from a sir_le elant.

1’“)
j\

’i"?

' I

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1 ,

germinstj n was mainly intlrenced 5y factors freaent in the raternel
tissues of the seedbail. Therefore, for the gurpose 0? this stuby

.C‘
L

germination will be considered to be a {y ical maternal

Characteristic.

EYjerigent R

 

Seeds from 51% plants of the broad-case, multigerm suivr beet

“"0 ‘V
!

varietv, U3 L01, vere harvested in 1957. In the fall of 1957, invoer

(‘f‘
(‘i‘
"1)

es .c tne seedlots inoiVifiuailg Tor sneed of (erwinetion, usinfi 20

a.

edhalls per seedlot. Of the $18 seedlots, 50 seedlots with :00d

’0
CD

(1"

otal rerminetion canaCity were randomly selected for further studies.

.edballs taken at random from each of the 50 seedlots

[1

Ten 3

f

(which will be desi;n ted as "half-sin families") were planted in pots
in the greenhouse in September 1958. Plants were allowed to grow for
three month, after which they were phototherually inducted for 10

weeks at 500 F (100 C) mean tenyerature and under continuous light
(TASKILL 1952) in a plastic induction charoer (fiQCfifiQAK 1960) in the
greenhouse. After induction the plants Vere left for host—induction
(ooOF mean temperature, continuous light) in the same chamber, until
they were transplanted to the field. All 600 plants of the GO half—sib
fanilies showed sooi growth and vi;or. fiefore trarsnlwntin; them to the
field, the rlants were distributed accordin; to their relative size

into two equal frours of five n1=nts, each group heir” ilanted as a
sefarate lield replication. The first rooiiCn: on was nlvnied on lay i,
the second on Key 6, 1959. Lithin a field replication the plints w’re
distributed at rendon, hoin: planted in rows 3 feet (90 cml a‘art, and

spaced 2 feet (60 ca) afart within the rev.

.éll— Plé:ntsa erminecl e*%7elflern; s;aeistxil}; a;yi exaed lieinalrt;uvrwt.

,-

./

./
\‘ r.‘ ,1 re [-9 .‘Vr7 1 1 _‘ )‘(Af-ZTV‘ - { b T " \‘F ‘} r I,'~~(?‘.\,"‘ ‘;“n :)- T,“ "_‘.} K I‘ J ‘39
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(13;:C-4T" :"l. i CV31" 'IF ‘ , THE" "1C! ( I". f I 11"“ rll ‘ 11.), ll" v «k. U-

N . .
ve t. rerve 1r wa~ vane 1. a r ‘i ,3: *ne :e ri J e ~ '_ e
r . v r “ l“ ,‘
between Jil , l and A l-t 9.
. . a ‘ ‘1‘. ‘ ‘ ,— r 4"
After dr'nri Tires"'~ ciearin , uv' Ll” or :1.) s . _. 's

)

‘ - . . 1 .3- -- ~ ‘ 7 . " ‘ I ~f‘
the n rcent of 113; s~ (cl n ii~ n:.<' 1h-) (a e .nwle 1L). ) r“ '1
' ' ‘»"‘ ‘ e, . #1,. N . .. r». e « .,. "‘, '1
terrinrtion for a 1 h-U (‘0 A ‘) seed..i. .1 ta ted in 1e .a-e tai~
f "\ ' I \
o? 19‘~. To fGCUT: unlior C‘Q’lll” , in s,~d‘-ii‘ vere lrlih,:ed

7"
\

in a tkinerature control CHWnUQT at a constant t“. erasure o.

1 .. .1. .. ' , 7: ,1, u 1 , 1-,' ‘ _ -. .r. . ( 7.
}ron the terminatlor ddkd 01 the EXCslluéflt it Man chtlle to

calculate heritabilirv e‘tiwates for steed of 'ernin:: on in tio ditfer—
r. .‘x . ‘ \ ‘~ : a w -. I ‘ . T7 - ~ 0' h . ‘ ~. ~ 7 . ' .‘s t ‘rv . “ ., - fl
ent WdYQ, i. e. irOm h:1.-le fanlJ MUmnS and from yi<nts vitnin nrlf-

Sib fawilies. Variance components were e‘tinated, in the first case,

from the f8fiil;’”wxfil s uares and fro: the Jigilies X field TQ‘JlCutdeg
mean squares; in the second case, from the plants within fmnilies wi
field renlicativns wean squares add from the n.ants X germination
replications mean Sonarefi (Tanle 3). Dififerences, si nificent at tge
1% level, were obtained between Families, and hetreen plants within

finil'es within field ronlicztio s (”able ”).

(‘4‘

The ., aes;mir<

fir]

enetic and environmental component: of varienc

I

 

*The term "percent of lost seedcens" refers to the rercent of seedpalls
in a seedlot that had at least one of their seedcaos (coverin~ tflu true
seed) missing.

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perfect cross nollinsticn in the ori ins] henul~tior, may he e3tius‘ed

U)
O O

( ATHER 19L9, cccraargv 1956) as foiicw

_ - .. . . Q /
setween hall-510 family variance (fififi = 1/8 D
' r
Within helf-sib family variance (05; = 3/3 D + l/h H

II

Families X field replications variance (6%) E

II
L1:
\0 o
5-?
:3”
(D
’1
(D

Plants x germination renlications variance (6%)

D is the additive genetic variance

H is the doninance variance

E1 is the environmental variance for family means
32 is the environhental variance for infiividusl plants

8 follows (Table 3):

g:

The variance component 5%‘was estimgted

ween Families }.5.) ~ (Families X Field nenlichtions M.S.) -

 

at
II
A
-L.
’D
(f-

r
where r is the number of field replications.

The variance comfionent 53 was estimated as follows:

6; = (Between Plants within Fanilies n.5,) - Families x Germ. hens, v.5.)
W o

 

n
where n is the number of germination reelications.
The environwental variance for family means (6%) was estimated
(FHE end Hoaxes 1955, JUGI 3956) as follows:

62 = Families x Field heplicetions p,s,)
1
l"

 

The environmental variance for individual plants within fauilies

t...

(6%) was estimated as fol“ows:

 

 

‘Q—v-u—

q I O — n o n 0 I ‘u ' I
6s = (Plants Vlthlh Farilies X rerr1nnt:cn renlicscicrs v.3.l

n

' ,c? - l «,i ' "rn‘lr. . .r~ ’.--x,‘,
The heritalilmt? Fit .0 Tor mama's-n: half—flu Trw‘ :' mean:- 5- _.z:l_e

'1

A) was estimated bv means of the follovin_ ferrule:

 

”'1; Le
a»

9 +
61
The heritability ratio for plants vithin half—sio families

(Table 5) was estimated using the follouini fa male:

h; = 6f?
a 2 + f
‘2- {J

Heritabilitv values are also given in relation to

U

 

L.

phd original
randomly interbreeding population (U3 L01), using both between and hith-
in family variances to estimate genetic variance components, and assuming
that linkafie and epistasis do not play an important role. In this case,
heritability ratios may be estimated from between family variances as

l owi°
f0 1‘3‘5. I)

N

1/2 D h r
h z: ::: *7

l/2D+E L5§+5§

 

 

0'

and from within family variances as follows:

h2 = 1"»? D + 1/5 H 5/3 6%

b -. c. -- a
11/1,. L) + 1/2 ['1 + 1.5 /, l...
A! 1 El"! + E

H

 

 

The hz values estimated on the basis of outween farily variances

correspond to heritability in the narrow serse; vnile h vaiucs Estimated

on the basis of within family variances correspond to heritability in lhe

E, 1/9 D, and heritability percenta;es for bk” 2—, f—, ani S-oay

I

fennination data, estimated by using halt-sib family variances, are {iven

c1 V)? "i
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n . >7 ,7 /" 1" _ fl ,_ ,
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;enwina inn 5.59 12.15 68.5 99.8

 

$1.

is the estimate of the environmental variance

L15

** 1/8 D is the estimate of the additive—genetic variance

#

x t
‘34“

2 . , . . . ,
ha is the heritability percentage calculated for oetkeen

half—sib families, according to the formula (1/8 D)/(l/5 H + E)
h is the heritability percentage calculated for the orikihal

randomly interbreeding population (35 qu) according to the

formula (1/2 D)/(i/2 D + E)

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2-day mination

a m

'e

P

3-day rernination l.L9 10. l 87.A 90.2

S-dev fenwhrfiion 1.23 6.15 83.3 87.0

 

’3

* E is the estimate Oi environ ental variance

lo nt estinete of additive and dominance

I
1
U

FJ.

ex (3/3 D + 1/L h) is the
variance

’3 . . . . a
h; is the heritability rercertage (in the broad sense) calcu—

lated for within half-sic families, according to the forvula
(3/9 D + l/L'r H) / (3/9 D + i/A H + 1.5)

if h‘ is the heritaoility percentage (in the broad sense) calcu—
lated for the orisinal randomly interbreeding population

(U3 A01), accordinf to the fornula (1/2 c + 1/2 H) /

(1/2 t).+ 1/3 l{-+ a)

in Table A. E, (3/3 D + l/L H), and heritability percentages, estimated
by using Mithin half-eib family variances, are presertei in Table 5.
To estimate heritabiliuy using the parent—offsyrin; re ationenip,
the mean genninetion values of the c0 (maternal) half-sib families in
1959 were correlated with the germination values of the so mother plants,
from which the half-sib families originated (Table 6). These parent-
offsrring correlations correspond to parent-offSpring regressions ex-
pressed in standard deviation units (FiJY and danish 1955). Assuming
perfect cross pollination in the original population, heritaoility
estimates may he obtained for the original US #01 population by multi—

timated r values by two (Table 6). These heritability

(I)

plying the 8
values may contain a certain anount of epistatic variance (CUCnbnndh

1956), out the; will not contain an; dominance variance (LUiH 19L3,

‘7‘; KW." ,. / “N '1 a" T' (",r‘vfir‘fl .-v\ f‘ 1 ' P4”.— v . .—‘\‘ ... -.‘<1 1 I“ - T" -. ‘u- -. .. 1.7.1 .
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(}E¢TIBLIV1C§YLL¥PA 1331L3 rw) Ftui}fl it} t’refi [XS L;14

r _ . o ‘ P _. / S , C ‘1‘”. ’
4—de¢ germination t.n«5m~ 9;.0

Q1.-..-": ’/'\.‘y."_ 'A
,-osy iaruinetton Q.+Mirn 9!.e

5-day ferminrtion O.2%7* €?.L

 

”Heritaoility For the ori ins? repiofiiy interbreefiin; yepUTeeiC.
vr-q , " _ 3 '1 ' . , ° ., ‘ r ‘. R V . ‘r
(L; A01) CQJCUIBted my uniti3lgin_ tne r ve.ue~ or 4, and ei—

u -. v .Lr'i. - - 'r-g «L:- .
preSSiNg neri-e»i!ity in “”TCP’ud e.

.‘

EYfleriwert L

 

In Aeril lQQR, controllefl cr sees w,re vane in tue greenhwuse

n.

1.

between certain olants from the surar beeu v riety, t3 hGl, which pro—

Huced either extremely *low or extrexelv ra{ii (enrinatin; seed in l§57.

'u

This was Fessiole of urin the original seeiehroducin“ riots ozonf Kith

{ome clones oi

I

which Envfier had crepe etcd in the green—
house.

'- “x N ' ._. Y's.“ ' .‘ ,,, .1 1 17 . ‘N I 1 . -h . r' .1 i- I .
;he plants Vere yflotfilnergatlg in1ucteu ?n tLe Minter of ij,/—

‘g.

1959 and nrofiuced seed talks in early Sfirin;. ”he plants were paired

\

accorc'nw to their ferminetion characteristics and their h; ocotcl
color. Contrcl of pollxo~tion was accomplished by gflmhs of white fiejer

l

'T. “he olents with the rapifi Lergination characteristic (3:5,

p
I

crva.t lwz

r‘.‘

5L7) had dominant-red hfjocotyle, and the plants Viqh the Slow gernini-

tion characteristic (:19, :57) tad rece33ive-vreen hgnocotfls. hence
the hybrid plants (?l% X 5A? and 357 x 5:3) could on iiemti‘ied by

hypocotyl color. After the seed obtained frou the plants witn green

1 , " -4\ —~ . r ' ’4 ‘1‘ , r- » u . ‘7‘ '-.- ' /‘ ~- « J* v. ~- I
L3 was ewrhlniteu, c e Seedling; h:th {reen hJ@OCO.;l: vere

tion. Leciprocal crosses (where the hOchS“ nlart had the re id iermin—

ation characteristic) could not be periormed, cec u e no plint: vere

‘ . ,- - u a- —. .~~ . .~ J- . 1“". ~ ‘ r' I ~ 1' Vv',~«' r-
hevbifi: ;lrmw seed «er IWU_J hl armiri vir1—'red Rd “Jeoc.i.

W L_." c.
‘ §

available

a .
The true hvhrie seedlirve here Viented in se irate “fit" in June

‘f‘ 7‘ 1‘ ‘ ‘5 r‘ P‘ ' . ' g" 5, V " “I" L r ‘np, 1 "' . I ’\"‘ . '
gene c the e eeew~in,s M813 gnototaermdllj lFLUC-mu in an

attemot to firefluce seeini rte oecacreas usherefions. ”fivever, ute
~ A L

A . 4 ‘, : . :3 2’f‘ ‘1 _ j, , 4 , ,_ _ ‘
croxeee h3vi> :I. :Vfiagoer l , ll-d :3 quydice 4?} V" u/ta seed.

'_"\ a x_ ‘\ 1", .- ~v‘ _’ ,‘f : .- ‘- 0 V A ” '.---~ 1 ‘ ‘ i . ‘- . ,‘
“fl” (“H.Cfl 0* T.'](‘{ 1.911.. .~ 4 .1.-..'Iv—:,11. u-l: 1'7”; "$.Jv .‘xt‘ij .« ‘ . L196- ill"! tllfi‘

cro‘fefl, clones or vefietative cuttinffi were a de from the seedetele

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The clones and the regaininf hybrid plants were iniucted nh~:to-

(‘+

r‘.‘ V "‘ + r r. . ‘ r . 1r I F~ 1 r~ N
herm1ilr alon3 wi h the a? nlant o- a rgnln and A? plant; a; a Q cw

ye- inatin polycrose pa ul tjon (fix1erinent J) in the Minter of 195%—

K

1959. The Y“..lr"-'r‘:ts were trenshlantel in the field on he? 7; 19%9.
The field exveriuent we: erren ed re a modified rindomized

connlete block desifn (Pflh3*3 10““ , in that twa entries (”slow" and

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entries (the four clonal nonulatiore) appeared only once in each clock.

All plow ' (eyetriee and nunltcate entries) ere di triuuted at random in

from which onlv the eeed of

fermirrfi ion exoerimentc.

The plants were planted in rows 3 feet (99 cm) apart end were

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snaced 2 f et (00 or W) art within the row. fairly uniform eeedstalk

develooment and bloominfi was ontained far the different royul: ions.
Seedhalls were harvestei it ennrexieetelf the some stage of ripeness,

between July 19 anu Au net 5.

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r drvwn', threshinr cleanin; S’Zinfi o: the seedtalls,

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3-day pertinetion data end for ci“Verent wocuiufion: are j1e+e i"

‘ ‘QTT f4

Tables 7, 9, 9rd 9. The fvur clomal cmrniwtiwne crevioed an exéen

l

{Beatinme of fxxe SYRILVCY‘“GHCQW.‘Varifflvlriisf, wirwze rd RAYS vr?th1r1:1 cirnie

7 ,.

are {ewetically iienficai uni car contain no _geefiic v riabiiitv. eince

'I

the between biant variance? 0* the fiur clone] Varnia‘iene were simiiar:

4'.

:‘i ta? hikivI/5e éa *‘HKETYii exit s afiie zrzr +1111 er“fiYV7me30fiil

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variance in -his exoeriment.

The genetic variancee For the sevre‘htini bflfluletione (T9918 7,
3 and 9) were oh‘a?ned my ynctrnctirg the pooled eovironiental vxriamce
(estimateo f"om the total v fierce be ween clonal plante) fror the total

variance of the segregatinu oopuletiong. r7 ccmfaring genetic uvo total

7.-

variqncee, heriteiilitf estiwates (in the broad cenee) we; be obtained

as follows:

h2

”9791— 3 w‘n ere
6%
6% is the total oheerved phenotyric variewce,
Gé'ie the genetic variance, estimated 5rom the difference between the
total phenotypic variance and the environrentai variance.
Heritauility values caicuiated according to this forxula ard ex—
pressed in percent are shown in Table 7, %, and 9.

1

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were: OT 9.1.x 714 Ant: . 3081,13 wxr'a. 0'.) .3! {1(2) flu . T. : ft—n.‘ * f >1” eel t) aer'm-
- e L,

nation as described in fix eri~¢1t C.

:- ,- , .' .. ;,. -r. ~ ’ ‘. .L..- . “-, WV» / .
inedn .eT'l".]n«~th.-‘rl ’le'lleb IO? tfle Li") SOLSCLC'L P'UUlllrsthOA: \‘qbl'uin

k

a 1 o !\ r1 \ ' ‘ 3 '. v ' - .Q ,~‘ ~ . . , M~ 11 ‘ ‘ ’~ ‘7‘ ,
and "reolo" frouoe 1n lvE/) dro tor the are GRJGQ oerwveq lrcm the;e two

I

populations or rancom ihterpol'ivetion viihin the tww areuye {eclycroes

froseni=s in 1959) are ebowr in Teule ll. The differences in percett

e . -. r V w l a H1 V ' I“ ~'.r 7‘ . I“ H ‘ or ~V\.r'~ . ‘ [a 7‘ '3 Y. : \ 5-. \ r w
germlnetjon oetdeea toe glee ann xa_1u ,“th3 1n 1%,? duh 1949 are algo

given. Since seLecfiion was Dr cficed in opfoeite virectlone one~half

of the pernihatior differerce oetweeh the two election yroune in ler

I

U)

will correspond to the selection differential: ”i”; and one-half of the
difference in fiercent garmjratlon netweei the two populations in 1959

r. - " r‘I $‘»‘\ ‘, —~ —..\.w« — a A‘ . "fl‘I' V'~ \‘r . F ‘V .
hill corresyond pm the gala fFDM oleCTlOU: v , assumlng no a Jnmetrlcal
selecticm response. Since AG = i X h2 (L‘3.::-.~;.1 l?

395, heritaaillzg ratios

1
1 ‘ I
I;

new be estimated in the follomin; indirect hey: h41= [Ml/i. fichever,

in this exoerimehb, he mother UlHHTS, wnicn were allowed to inher—

oollihete wihhirlifiue‘tma groups, orijineted frou a reryxeLv inter-

breeding population. Thus the eweecied gain fro; saleciion can be only

0 .

one—hell o

’4

that, vhich could be expected in a self-pollinatin: C“??,

,4.

r

. . . ‘ ‘1 ' ' \
or 1n crooe where poll1natlon from both parents can be contrOlled (earn).
Therefore, to obtain heritabiliiy eetlwu‘es for a CTOSS‘pOllihfitlflg

crOP; Wmich has natural barriers to self fertilizerion emf controlled

crossing (1. e. suhar beets), the rein obtaihoi from select10h VAC)

1

1‘
[J-

\_z

{n

has to be wul'inlied by tw; and civioed by the seleczicn differertj

<r‘

“ l
.’.u
;

fellon

T:
5'". )

1
”1" u r g
- .' H ‘1 ‘1.

X 2 2413

 

-J

) ”Ere eftir fed for the aneraior
F'I}.

.weee values were n'ed to

5-1 (-p arr: 1 rm 3* n o Tamr'mul H f‘fir o t" 1..
‘ a l ' - 4 u A. , . _L._ _ ., J A .‘

v x

..J

eat ( 1C 0, lupin).

PJ.

f‘Jl’TWQT‘ITiEQY'C t,

 

Ct

To determine he influence 0? environnent in fiiFTer;

\U

rt years
on speeo of Lerminetion, and to disclose thether gerefintll, different
plants react in the sane way to the oiiferin; environment of the tee
years, clones of the 23 plants, from wnicn seed was ooteined in l957,

were P anteo in the field for seed production in 1039.

/ .
The nlents were nroneweted veretctivelv in the treenhotse for
. ‘ _ ~ (I '__l

[I

Z,r o- l9€?—19>9, and treleplanted to the

(D
1'14
p.
:3
r +
*3-
1D
<‘
,._.I
"J

1% month, inouct

field on bay 7, 1959. The nlant were distr outed at random in one

field location. The seedoells, 31”“: With the remnant seedoeALB from

('7’

he genetically identicel nlente, were ferminlteo simultaneously 7n a

/"

\

1ahoretorv erneriment. Two germination replications, each consisting

O
“a
h)
C)
(,0
(\

edhalls, Here used for each seedlot. The germinetlon Cate,

V

e2nressed in nercent germination, are listen in Table 12. The analysis

1:.) 114'
l... 4.2.

of variance oeta are given in Ten

Although the germination of the seec from the 23 nlents differed

.L

Signi?ic:ntly in the two years, the plants did not react oitferortly L0

the environnents c; the two genre. This can be seen from the non-

...

signiiicent Years X Plants interactions (Table 1;).

(_ W "Y ‘ 1 A I“ " rr‘! ' f‘ —-\v n 1‘ r «1 ‘ v1 r 11 ‘ ‘ r v - . \ -—~
I I 4 l I ‘ \ I r 1 I ‘
- 1 ‘. - a. l .A. J - - l I ./ '
x- . < ~ » r“ 7‘ ‘ ‘ -- ~ , .— r —v- , .
1-‘TI ‘\ ‘f ‘ ‘V ‘ . I} '7'" f N I ‘ \ Y ‘ ‘ ,. ' ‘T “ I“ F ‘
.’ \ A I 4 l. . ‘_ ‘ , , t i - t _) I 1 -t
f "‘ 'Y ‘ I] ‘v rn "W\
‘ 4' f 4 ‘ ~ :1 fl.
\ ‘- I
A ‘ 1
.’-|‘" 1.1T: - *I‘J(i"r Cmf“ “'
A-_ -' .1- .r ‘ j 1 J
' 1.
FY \' , . ' A l 0 A ., a ‘I L 4 7‘ " ‘ . ‘_ .7 "‘ —
fit "I" "vvréywry 1V.",‘ A‘ ‘. {NY} Tr‘d,'-rr 11",», I Y‘ {7 9V": ‘Ert‘ ‘ if
v ,- 7“ .r\‘ r~ . H /, ~~ \ , 'r 4" ’7 \
3‘!“ 1hr... Lmu 105.. “twig 1|(L7 1w”,
' ‘ - ' ."I ‘ I .’ I x‘ I - .“, -‘ I .’: ' l , ."

 

:~.
K.‘
b
Ks.)

O
'._) \J

C

.. “I

l, "4‘ f". 1‘ r7 {j
1' .
f

V0 (”Q r-‘
3

L, ,
’- rx - ,
0 If ‘ j A C) '7 "I

:>
if) \u \n O I-

’ /
"\ ’\ (‘N (r r\‘ 1 f‘ i".
A ‘ I r . . ‘
/ J v e - ‘ ‘X ‘ SJ
/ , ." A , \ A _3
CI (5; ’5 £3 ' pi * r I:
/ -I ' / r‘v '\ "\
h _; .1 :2 "j ‘3 t O. H [:I
. P, — I, .
'7 (I, '75 Ag“, r7 , r, L.) Ir)",
‘ V" V . ,1 . 4 ,/
9 L} ’1‘ '7 :1. C4 "4 (- f," O f .1 11-?
J ' x J A - _
Q A: '7'} CI.) 1(th (:2‘ .: ..
'1 7’ ’ V ‘ I a j
10 GE 7? c0 ec an (m
‘ V ’ I I, ‘ 4 k
‘1 1 0:.“ "f 1 C rs r. L; (“IQ ’1 c)
.... 4L . I , 7. x r
I? do fit 0‘ “5 MW 00
V I -A - / ~ / r
" (5 O ’7' "‘ Q i.‘ L‘ ( J Q C
‘1 / I 5 .‘ ‘ V I — 9 )
(VA n i 19"“ ,r.‘ I”) ‘ _“ C "\
11" /‘U 1' 4’ 7 1" " v f '. v
1 a o <7 c. .. :~ a
’ I . _ ,’ ,J’
A Q t. C / C L. ()O (.11) 4:},

-q
o
J"!
{‘1
J‘t
“C .
J7
..J
1‘

.1 an
._J

W,
'_‘;

.-—J_J»——J...J
O}:

O ,

e n, .,
‘0 J

n .2
r§~o
OJ)
O~O

or D

..J

h '

‘r‘

1,

Q
n
w
.A
I
V‘ .

“Q

.._J' .J
A I

,.

J
‘x
’I
_J

_: "\

\ < .4
.2‘
1"
V

I‘\

L. 1

(J

‘I

1%

L._J

\

l

J J
....J

. "\
’ \

\_J

,_J

c J

..J
6*

”K.) I\) Ix) I\)
\k) [\\) v—J {A

a" Q" (I' 0'? (‘2? 1 )

l / ' ,' I

Or“) Q: of} P C ( (x ()r)
A- I ,I I I

fiverexe C” 211

4- ~~ ,1 4- r‘ e ”a“ '7 oo 7
VIIQTHH.) ’ /\.Q [#1.] 7 1 I) C QC .‘ .,

LSD Cf hetween

T‘ 'ir‘ts 11.9 :11...“ ‘lkx.g' 10.5, 11“), 11“,;
" rt'.’
1.?! 1 ,r) h0+"~'eely~l

arts 1?}? 16.7 1“» i 15*]. «(.1

'3
..1

CD

..L’
‘5.-
U)?
r\)

o

m

4‘} f/ j
4.. n, ~01}.
. rr.’
1.39 1 e het‘xreen
~ F' ’3 r'\ p T A
7' '- 2‘ 4 7'
p {r J . / ‘1 . .4 I . ...

 

*~-*-—.~v——- ...—..- -..... “...--... —.._—.-._——.——

 

-: :(‘3‘3

r4.

flLD P

5

X UK

)Al+

T .
.L

E).

" (O
I'!' L

IEJIL;

1 W ‘
ti

(U

«VS.

J

Y

\
I

Tm”

)
1

U3

4

Jenninqtior

(...

7-d.7i;.*
O

f
4
/

['7]

3 O U H C

 

/03:]-

l .

. ‘5
\V.__41((::

l

v

\ T1
\Y‘~

1.2)]—

\0 LI“. .Y\

\o p- ,4
O O 0
LA -

.¢

I
q.
H
=J\ (\j
0 O

(\4 (\I

.o
L
1.0h

3'; s»
J, 3:
[‘9‘ ..-.
T. - l O ) \O
\r J Li\ W
O O O
3 C13 r—{

O‘\ H

C‘\ (\— I’ \I

O O o

D- \T (\1
(\2 Ln

0) C J

3
Plants x Years

a
Years

“I

1.11

l.L2

ror (Remainder)

1
J-

1‘”

'-

91

 

“nificant at lj level

l

31.

*e

'7‘"? '- " - ‘ '- ,.,. f“ . 4 ,. . ~ .- ~.-- 7 .J' .- 7-‘ 4L 7' 'x'
"1? OC‘. 0’ + "! «Ta-”‘3‘ 0' 860%”: , .' "H BEG” ." 1.” ‘:‘ 1b}. ’T‘

-* ..- ..._. —.— c. .-*“< -----—-- *- ”...-v. —a-..- --—‘ w-—. ..v —_A-_..‘

r r, fl \ r‘ .1"\ c. ‘. 5. l ‘n . A - , 'v“ ‘

lit “)~oc— Q er! - een o1 .errin .Ien O; .ne “1 r nee- Jor. tn
I- ._ W 4. _ - c ,‘I .. ‘ 7 ’\ L _. ( _ 1.. -,.. --
. C o- -1- '~ eev‘ual - 3 cf f”; 1"] . .3 “T" 1.3? [lam ~” 7 r‘ I1'. ~-. '2.

7 ' ._ '. .J . . 'I_ _ ,‘ ,_ '.‘- ' _‘, ‘ m “A ,. -.J .L r ’ . _._ y: '. ,.
ODtl lT"-3;.i r."»‘-"\‘7"?'?rl tit"? U“..-";:f' C,‘ 1135.4, SBAICit- :3 air}. ..i'?) a" 12:7 ~,r‘..,.in:zT,10.Ia

m .' ._.'- - r.. 1_.L-
+ .he tern 'l®~3 seedc.':” vIlJ r~,er

\

1'? N'ler 1 «" '. «7 AVV' . . 21.». w . a. . N
‘iU L53 411+) . til -.I At. -- ll hm... .;~?8'~_z1.313 here 1.1.1. 2%».35 .J‘Q Farthirzd
. w... -.-. r 7 ,. ‘1 « , . -r-- J.' . J. A ,,., —: fl U ' . .
3'.- TO». 34+ 3.1., #.l‘;.’ .3 1. '3 V. . ~ J ‘- I]? “V'I‘CQI‘ OJ. .11.) (3+3 SCIEUC ;. 3.? {.314 Cl”? fdl'e.‘ .t

“+4 «~qu 73+ p. , '7
uw)gb Tet v, Huekgfi. ciolul;

’V fl _" _ _- 1' . ' ‘_‘ .‘ y ‘ _‘ . 1.. . .f ,. a I I

i --, = A? w'r? nee '11.hr; ~:-Lc 3'u~i“' (an :iluU' * , L; 1 d
m ‘ ~ ‘ ’ ‘ " ’ I r. u ‘ “a 1 " v f' ‘ “ u I ‘
.~ + three rem“ Atter tnzd ”In; i14> Maren»? o. 10 exe.r ;

I —( 0 4L 1' — l 4 L _ I
' ' I‘) I f.) A!— \ . y ‘. O 4 \ ‘1'
]”‘7 +n9 CflerFilh‘lfl V lflm lCT tr ; F~ Maze 1 . l”? in f L:-
l
\l ' . ' ’fi .-' , ' '. ,. “fi ., . - ' . -
r'1 “1 13 e. . I ~ 11 'c t - L ~~ i:1 iWH' - L‘ " 2 1:1 ‘J.”Ic€ k)f

 

I v x _ _ _‘ j o W ‘, fi - ~r ,‘ ‘ \ a _‘ q- . ~ ‘ ‘ N '7
one of the seenc-ra (51a. c verim ,xe +rile 2"4. 1; n.--ir .

1'7"?

 

. 3 1 ' 7 . ' " «I "w . ' . '1 t .' "h" '1 ,. . - ‘ .
’L' ..L. ' Jr.“ - .' _. ,‘ x 1 . .gll. I ’ IT. 1" H "-v 1%, . O ‘1 '1), ill) ‘l t.‘(‘: I"
~ 4. '1 ,.L ' - -‘ ‘ r J ,. . ..1 , fl ' j 'r'4 ‘ .3 fl‘ ... y, A . \ , 1-, .. .‘2
c *re,,-3~n n1' er I m en a; +0.1- , n'v'w P”IF:» ..., to 4 her; .
I'l'L ? w~ r‘ 3- a « ~ .--: -- '~' ~. . .m; yr"- :
ui-i-3 “.~-..-€:'x ,. '>"c«. . . ‘c c. I .-u: -~- ivhc—.--n I'M‘
A“ ., ,f ,4. 1 “h (“mgr-:13 (,5 3...: .- ‘, ”I (.. 4 ,. wava 4 A... , - ,n y, .3 -. . -' We
L._.IT'r.1-.;, I-.. -. - _ .. r.-I-., q“: -1 .- . :. . .- Ma.) 1““ 1 - cut
"I ' " ~‘r r- ~ » -r \ 1 a 1 I
+w'4Y‘:O,~‘”r "-._., x”) . .. \' /II’"I 3_'| 7. '\ (‘W‘flv‘D .1 ’0'. (1, QL '14f‘_-CA"L
J4 . L , . I- ._\3 _ I I 0 'X 4 - . ._ . ‘1- -.. . .. .a. -.. . .
_".'L
A /"“" r - +fi.v ‘1 ' 1(f«‘ ~$v A t I] r‘f ‘ .._. 3 v '
A '_" '2 ~. I‘ “ ,.‘_ r"- .3 .\
+..‘. 54 1; JV L _ 4 t’-‘ . 31"} L/ 'J J i . L’ 3r. \r' u ~._ ., J . eu'tC‘ .‘»J 1 . k t -_
.' . o H a A- _ - _ ‘ . u ;, Lc ,' 11
,- ~. -, ..- 1 ,~ . + . 7 . - - . .. . , - «
(TY. ivifl’] f it - ._ L'r Jere, J -' 'F "J " l e.( ‘ 1'4“. ‘1
1/3).
'u'l» fi \ fl “(‘1 9 ‘ 1.x”) ‘ v W‘ ‘ p r I I "3. W )1", 1 o ‘ 'N - 0 fix ‘ L ‘ f a
L -;-", i :.H.qq -- (5,0, Vlriu n P" _.' * "uulC’r‘Dii ' ( -..” ~* .

ti htnecn ot +he :eedcene i? sweeticeil' cwr‘rol‘e". . » 'He 'ercer
g. t. .
o e ‘9”i‘ . .1‘11“ 1f. . II "5” x () v.1. ‘fi - C' A, (i '\ I: r _~ i». -,1. ‘J. .1 vulj VP 17‘ ‘: 'r L. : -
Q. u€.,l U4- 1?, ’-.Il(‘..l 1.. f3 :z.._..-T1’ dip/MAW :) 4.“) I“;-.'.-'..,x\l‘\"'~h -.~ .7.) v .I.1
-, , -.--.L' -. ‘-" .. i , ' -. L ‘ ,- ., ' . .—. .
ciir«c r* 10 . it Ir~c l in :c -e r -li ar in *Inn o bra..l.

ner e? eriaerfe are re mired in deterxine the ;*ec151c enviror-

yu
‘r‘d

* 1 r 9 —. 9 ~ * . '-
r? . ‘ IJI' . P . (J .1 k " 3 ",‘1 ‘ Tl ) Y“ L J: 1, ‘ 7.? ‘ 9 “ 1 ‘
v.9 ‘ ~ -,—~ - - {r 7‘
I 1 n . J ‘ L; I ‘ I 'J . " ‘ 9 J ' 11.}
1;” v I‘;',"7 1‘" ( [C
I; \] I ‘1 p,» L, 1 .
" ~-s 1 ‘ x
f a ‘ i Q 5 7 Y I a 1 l 9 S O
n *-
r . ’3.- L t “.0.
W‘ a ‘ ~~‘I
k—l 4 lI‘VE) .4 Jr S PJ-’ (

 

clone 7:9 sfimw+) 1 0.3 12

Clone 3C7 ( 10k) 1 0.3 1

(\"

k}

Clcne 5:9 (rqfi1++) 1 10.5 1
CYOHe 5A? (9 rid) 1 7;.5 17

i " Q‘ 1‘: S‘117 h'f‘x')?fli d 2/4

‘1

Slow seWection Erfiue Q C.] Progenfiei 0' 910“ i””UT A“

Hamid 99 elect Ion rrnuh Q 2“.Q Freneniea of r9“id grape LR
Flanfe Uiih 999492119 R nrnveniee of each

th5W? nfi JQSt $99333r3 4.? Vlgfit Havific F0 lnSt 17A
399(3ch755 1n 19L)?

4‘)

a\
[h
_)

A)

Fleets with eeedh9713 9 hr 0:395 Cf each

havin at 1929* 105 7? ”0.5 9119: anih at O‘St 1‘6
loet seedcefis 13, lost seedcaws

in 19 5"?

 

Cfirrelation betveel the ha: 99* o: Correlaifion Dciweew *ne nu we? of

lost seeflcqns and the 2~dnv 105, seedcens and the 2—09
99“"1n4+1 n counfs on the basis yermihawizn courts, on t
of SO F1?ntW(1957) : of GO fa¢ily mean: (1959

V :3
0

r = 0.67“”' r = c.60'

 

+ "310w" r9f rs to plaeps producing 9x
++"ra:id" refers t3 plwnts graduein; ev+
**Bignific9nt at the 1; Jevel

:SWOVW5’vzerng._.,ln":v:

93:
e 61! rajidig ;”rmip;tjn;

,‘ 1‘- _‘..'“"“T
DI f jug/31.21?”

"I
-
‘-

djffererces DGVEBLW%?1'HT56FV€d i'1 :9eed.o‘

. LCA _.‘

:4
m
.1
(D

-.

of sugar beet seedlfits :arveeted frsm iniividmai yldhi . The engeri-
rental results (EXpérimefit A) re~confbrm previous Deserv.bi9n5 tndt

r to the characteristics of Lre raterrnl
tissues Cf the seedbell, SUTT”LCCihf ;ne true seed . 99th wn*3ica] and

c.9nical attribufiee of the seeficall have been Leeonetrnted to irfiueuce

rate and precen: germjration. Aiihouft 5t 19 veesiole tuLt 9999 dif—

'_r 'r A

ferewces may exnst netween rhe g‘rwivntrnd 0n *ne 1rue s~enb, fine e

f e eed cf

3‘
’25
If:
H I
<1
<4
,3.)
xi?
J
.21
fl
,4;
1:1
(L;
(.9-
...4
a:
w
(E
D‘V
..J

A marked positive relr

‘I ~ A ': 4‘ '. ‘ . 7.7 \ I [‘ r‘ '4". r 1 ‘,"\ 1‘ 1 'r' '7 m r\ r L . r2 . .— 1 ‘
””rmlnntton 831 tlkf.“€:¢ O; SiedCst, wflex fin; Ln-b€? WQS LBJSHFHQ Dy

(

the percent of seedbelle having missing seedca33 faoxe 1A). ;nis
euifeets that the physical characteristics of the seedhnll he; {lay a
relatively important r319 ir cetgrwinir; ewwed 0: L~r inat an.

The reenlfie of the geretic eY erimenhs (x, C, and D) jive

definifiive evifierce that the jerwiretion rqfe of the sugar beet seed

is under renefiic ccnfrel. Th3 heritenilLig ratios (he), obueived by
different stwtieticnl technicnes, for the .—, 3-, and E-flay Lerxinatlcn

counts are surnarized in Table 15. [hey are coneihteniiy V'ry hi-h,

, ° , ._ w ’\ h ‘ “,9 U: .4, ,3 , -.. - r". _, M - ’4 ,._' ' 1.-
e peelallg fer tne 4—, “no 9-04y Lernwnatran CAane. Lne H”Plc belie;

. I 7 I .7 . «7 .I . . 7‘. . l\ x. 7 7. . II ..l I: 7. I. . . I I. I: I . I ,u \ V. .II. . y 3 x I . . . _ I .. ..7 .
I7 LA... L I + CrQC .«LQLCHF {7 ., .. ..I 3 .( C 7. 7r LIFO, L ..... .... um .10.. C. I: OCH CL Ic‘ 5.1, a! 7. 4 [.91).
. I ‘ .I I . . I O l —

I

9 1.. A . I; . €- .. \ . I \ I.I. \. \4 V .. .- \fll. . .JA . I ‘I I. 74 II! .I, .I. . i.. ‘ ,.. \. ‘1 4.«III I \...I.. ...r1 J . . I c. I I I. \I I 4
:43 .... .. .e.._......7_; ..2 It 7.7 .I “I.— AOP 7w 7, NIH QMIIWr.» x+LCQ.....CL "Pb )IL...» Q4152I+bQ 3PM T.,...,.r>_n¢.+f,.(. I... ..q.._.r+LC 0717 Z

(r

 

. \. I, \l 1: I i II .1 \
vk.‘ Ant-U i r v H. » {EPWVI
7 . II I . . . . 7. I .I... : I I. I 1 I .I I1 I 4} I .i ‘1) I .3 l . . I I a. 4 \ 2 ,. ....7 I .I. .1... I .,.) x.‘
.I..... r . . I.. In.) 7, ..7 _ , .. .If, I ., H...“ I: _. .7 ,. i .7, _. _L H ... .7 .
I+ ,I I... r .fl I 9% ;v I4. (\ IV .I. «I .7 | + L m7 0 (b r r WI I (I g (m ;.\ o- ‘e r (\- r: W. + w\ ”INF...‘ (:1F . . _ fv‘: . a P... ? AIM. /I\ r I4 f n F . bu \ g Ir J I\ (I m. P w. (£LC +— LIA-fl L» I. r .l r o. I I+r -/\7_ ¢
\ r ......I ,. I. III... 4) )I .... \l 7.: I1. , \1 Jl\ \} .1 J I; J: ,. . .. I ....In \. II... .II 4 4II \: \I 4. I1. \..., . .1... I \I ...v . . .. . .7 ,y
ofl.(~ 7.... rKI LN U. r -rC ..., .. ,. C1 ICFN fl CUI. .... YIN 5r L 7- +0 \,.-_ INK/I r 7. _. L MI. r. . .I..( I . _ .l . W. H) .I 1. L IHI rtH £2 CC ...I r 21:; G 7‘ ., DC, I LC(
4‘ r t + r .4 :4 v o 7
II\ : ‘ .l I l ..l... . I. .‘ a .. \;\. I .. . ‘ ..II ‘ QIJI .oI . ,u .9 . . "\I 4.. ...u «:11... III. «7I II. J u I I l _ \. Ix} . I 1:14. .7 .I x. A \ . I .1
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itahility eetirates, ontained for dinerent populations using

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