Plant Breedingl16, 337-340 (1997)

|c.;1997Blackwell Wissenschafts-Verlag.
Berlin
ISSN 0t19-9541

Inheritancein seedsizeof alfalfa: Quantitative analysisand responseto selection

R. Grunrcr and S. R. SnrrrsJnr
r DeutscheSaatveredelung.
WeissenburgerStr 5. D-59557 Lippstadt,Germany; r Departmentof Plant Science.University
of Manitoba. Winnipeg. Manitoba. Canada. R3T 2N2
With 2 rables
Ret'eiredOctoher2l , l996iAcr:eptedApril 24, 1997
Communicutedhy W. E. Weber

Abstract
Seedsizein alfalfa(Medic'aoo
satfuuL.) hasbeenpositivelycorrelated
with seedlingvigour itnd early growth.but thercwcrefew publishcd
reportson inheritance
or selection
f or this trait.The objective
of this
research
wasto estimate
components
of geneticvariance1orinheritirnce
of alfalfaseedsizeand determinethe mostefficientselection
method.
Components
of geneticvariancewereestimatecl
on seedand pollen
plantsof 'BIC-7-WH'andtheirprogenyarranged
in a North Carolina
Designll matingdesignundercontrolledenvironmental
conditions.
Threeselectionmethods,difl-eringin parentalcontrol and selection
pressure,
wereusedto determine
selection
response.
The seedparent
genotypehad a major role in determinin-q
alfalfa seedsize.but the
genotype
of the seedhad no influcnce.
pollenand
For genetic
studies,
seedparenteffectson seedsizeshouldbe measurecl
on seedharvestcd
from progenyplants.Seedsizewascontrolledby additivcand nonadditivecomponents
of genetic
variance.
Heritability
fbr seedsizewas
41.3%.Selection
for seedsizeu'aseffective
and a significant
shift lor
largerandsmallerseedvuas
attainedafieronecycleof selection.

the results. The strong seed parent effect and inconsistent influence of the pollen parent suggests that genetic expressron for
seed size may need to be measured on the seed developed on
the progeny plants.
T h e o b j e c t i v e o l ' t h i s r e s e z r r c hw a s t o e s t i m a t e t h e c o m p o n e n t s
of genetic variance involved in the inheritiince of alfalfa seed
size. In addition, three selection rnethods were applied or-ra
r e f - e r e n c ep o p u l a t i o n t o d e t e r m i n e t h e m o s t e f f i c i e n t s e l e c t i o n
method to increase alfalfa seed size.
Materials

and Methods
P l a n t m a t e r i a l s :T h e r e l e r e n c ep o p u l a t i o n l b r t h i s s t u d y w a s ' B I C - 7 WH' (Barnes et al. 1977). tr broad-based germplasm developed for
e x p c r i r n c n t apl u r p o s e sa t U S D A - A R S R e s e a r c hS t a t i o n si n B e l t s v i l l e ,
M a r v l a n d .a n d S t P a u l . M i n n e s o t a .U S A .

Cultivation: Growth-room conditions were l8 h light ( :400

n r E i m r . s ) 1h
6 d a r k . 2 5 C I t 3 C . I 5 3 5 o Ar e l a t i v eh u m i d i t y ( l i g h t r d a r k .
Key words: Medit'uttosutit'u heritability seed size selec- r e s p e c t i v e l y G
) . r e e n h o u s ec o n d i t i o n sw e r es e ta t c o n s t a n t 2 2 - t 2 C w i t h
tion methodology
a d d i t i o n a l l i g h t t o l T h w h e n n e e d c d .G r o w i n - r m
r edium wascomposed
o f s o i l , s a n d .p e a t a n d m e t r o m i x ( W . R . G r a s e& C o . L t d . . W i n n c ' p c g "
C a n a d a )( 2 : l : l : 1 b y v o l u r n e )i n 0 . 5 l m i l k - c a r t o n s P
. l a n t sw e r ei n i t i a l l y
The relationshipsbetweenalfalfa seedsize,seedlingvigour. and
f c r t i l i z e dw i t h 4 g N P K ( 1 0 : 4 8 : 0p) e r l l o f g r o w i n gm e d i u m .a n d l a t e r
subsequentforage yield have stimulzrtedconsiderableresearch w i t h a d d i t i o n a ls o l u b l cN P K ( 1 0 : 2 0 : 0* m i c r o n u t r i e n t s 5) m l , i l w a t e r
interest over the past 50 years. Black (1959) concluded that e v e r y 3 w e e k sw h i l e w a t e r i n g .T h r i p s a n d s p i d c r m i t c s w c r e c o n t r o l l c d
early growth and developmentwas relatedto seedsizefor most b y s p r a y i n g D e c i s r \ l ( A g r E v o C a n a d a I n c . . R e g i n a . C a n a r d a )o r
agriculturalplants.but harvestableyield often was not. A high T r u m p e t r M( N O R - A M C l h e m r c a l C l oW
. . i h n i n g t o n .D E . U S A ) o n c c p e r
correlation betweenseedsizeand seedlingvigour was reported w e e k . P l a n t sw e r e c u t a t 5 0 7 o b u d s t a g ea n d c r o s s i n gw a s d o n e o n t h e
s e c o n dr e g r o w t h t o e n s u r eb e t t e r f l o w e r i n g s y n c h r o n i z a t i o n A
. verage
in alfalfa (Beverid_ee
and Wilsie 1959),and the advantageof
g
e n e r a t i o nc v c l ei n c l u d i n g o n e r e g r o w t h w a s l 9 w e e k s .
largerseedincreasedwith seedingdepth (Erickson 1946).Other

researchershave suggestedonly a weak relationshipbetween

seedsizeand seedlingvigour (Nel and Burgers 1968),and that
the relationshipdirninishedwith plant age (Smith 1961).Carnahan (1963)found a positivecorrelationbetweenallalfa seed
weight and unifoliate leaf area. and unifoliate leaf area and
seedlin-s
height at 4weeks. but there was no direct correlation
betu'eenseedsize and seedlingheight. Despite the interestin
alfalta seedsize,there has been lintited researchon the genetic
control of this trait, and no reports on the selectionresponse
for seedsize.
There have been reports indicating a responseto selection
for seedsizein other herbagelegumes.In three cyclesof selection for lar-eeseedin birdsfoot treloil (Lotu.st'ornic'ulatus
L.)
the averagegain per cyclerangedbetween6%,and 20% (Draper
and Wilsie1965).
Previous genetic studies on alfalla seed size relied on
measurementstaken on seedharvestedfrom the seed parent
plant involved in the crossingdesign,which may harvebiased

u S .c o p l r i g hctl e a r a nfc' e n t ef r o d e
stateme0
n lt l:9

9541 l91

,1604

P o l l i n a t i o n :C o n t r o l l e d c r o s s i n gw a s p e r f b r m e d u s i n g v a c u u m e m a s c u l a t i o n . w i t h t l i e \ r a c l l u l nt u b e s t e r i l i z e dw i t h 9 6 7 o e t h a n o l b e t w e e n
t w o c r o s s i n g sP
. o l l e nw a s c o l l e c t e da n d a p p l i e dw i t h f b l d e d c a r d b o a r d .
Standard petal was partially removed.Open pollination was perforn-red
w i t h o u t c m a s c u l a t i o nu s i n g a f l a t t o o t h p i c k . S e l i p o l l i n a t i o n i n v o l v e d
g e n t l y r o l l i n g r a c c m c sb e t w c c nf i n g e r s .
Quantitative analysis:Twenty-lbur plants. three setsof four pollen and
four seedparents.were randornly selcctcdfiom an initial population of
'BIC-7-WH'and
a r r a n g e di n a N o r t h C a r o l i n a( N C ) D e s i g nI I c r o s s i n g
d e s i g n ( H a l l a u e r a n d M i r a n d a 1 9 8 8 )u n c l e rg r o w t h - r o o m c o n c l i t i o n s .
F o r e a c h c r o s s .t h r e e r a c e m e s( : 1 0 f l o w e r s )w e r e c r o s s - p o l l i n a t c da t
f i v e c r o s s i n gd a t e s .C r o s s i n gd a t e s w e r e u s e d a s r e p l i c a t i o n s .A l l s e e d
p r o d u c c db y c r o s s - p o l l i n a t i o n
w a s h a r v e s t e da n d i n d i v i d u a l l ym e a s u r e d
t h r o u g h a c o m p L l t e rd i g i t a l i m a g ea n a l y s i s( D I A ) s y s t e r nD
. igital inrage
p r o c e s s i n gs y s t e mu s e d f o r s e e dr l e a s u r e m c n t sw a s d e s i g n e di i r o u n d
a D T - 2 U 7 1 ( H S I ) t r u e - c o l o u r f r a m e - q r a b b e r( D a t a T r a n s l a t i o n I n c . .
M a r l b o r o . N { A . U S A ) . M e a s u r e m e n ts o f l w a r e w a s l r n a g e X ( D r L .
Lamari; Departmcnt of Plant Science. University' o1' Mzrnitoba.

0337 $ 14.00'/0

Grunrc and Slrtru

338

method. For both LS and SS populations, eight half-sib progeny from

e a c h s e l e c t e dp l a n t ( 1 2 x 8 : 9 6 p l a n t s p e r p o p u l a t i o n ) w e r e g r o w n
and randomly intercrossedwithin their respectivepopulation under
greenhouseconditions without emasculation.
The third selection method was a form of mass selection with no
control (NC) over the selectedparents and a selectionpressureof less
than 1% (basedon weight of size separates)for both large and small
'BIC-7-WH' was mechanically
seed. A representativeseed lot from
separatedusing a seriesof dockage screensof varying sizes(Seedburo
Equipment Co.. Chicago. IL. USA). Twenty-fiveplants from the largest
seed fraction (screensize > 1.8mm) and 25 plants from the smallest
seed fraction ( < 1.0mm) were grown under growth-room conditions.
Plants were randomly intercrossedwithrn respectivepopulations with-

Winnepeg, Canada). Seedsizewas measuredas the area of seedimage'

Previous research indicated that a large sample size is required to
eliminate seedsize variability resulting from the number and position
of: seedsin a pod. pods on a racemeand racemeson a plant (Gjuric et
a l . 1 9 9 3 ) .S a m p l e s i z e f o r t h i s r e s e a r c hw a s s e t a t a m i n i m u m o f 1 5 0
sgsds/plant.
Five full-sib progeny plants per cross,totalling 240 plants (3 sets x 4
females x 4 males : 48 crosses),were randomized in five replications,
with one full-sib from eachcrossper replication.All plants were crossed
to a single pollen source plant and also self-pollinated. Seed from
individual full-sib plants were harvestedand measuredthrough the DIA
for quantitative geneticanalysisfor seedsize.
The analysis of variance was calculated with mean squarespooled
over sets of parents as proposed for the N.C. Design Il (Hallauer and
Miranda 1988). Additive and diallelic variance, based on an autotetraploid model for a factorial design, were calculated according to

out emasculation.
The three selectionmethods produced the following six experimental
populations: (l) large seed full control (FC-LS). (2) small seed full
control (FC-SS),(3) large seedhalf control (HC-LS). (4) small seedhalf
control (HC-SS), (5) large seedno control (NC-LS) and (6) small seed
no control (NC-SS). A greenhouseexperimentto determinethe success
of the different selectionschemeswas establishedusing the six selected
populations and the unselectedreferencepopulation'BIC-7-WH'. The
experimental design for each population was a randomized complete
block design (RCBD), with nine replications and four plants per plot

Wricke and Weber(1986):

d e : 2 ( o i , , + o ? )- 2 i 3 x o i '

ill

oto:6xol,l

t2l

where o2o: additive genetic variance, o']o: diallelic (non-additive)

genetic variance, oi. : variance due to males. oi : variance due to
f e m a l e s ,a n d o i p : v a r i a n c ed u e t o m a l e s x f e m a l e s .N o e p r s t a s i sa. n d
no tri- or tetra-alleliceffect) were assumed.Narrow senseheritability
for seedsizewas calculatedas:
h' : (oto* li3 x 4) (o* + o; + oi + o?)

(36 total plant per population). The plants were randomly intercrossed
without emasculationwithin each respectivepopulation. Seedwas harvested and samples containing a minimum 300 seedsper plot were
measuredthrough DIA for seedsize.Seedweight was determinedon a
seed sample of 150 seeds per plot. Mean separation analysis used

t3l

q'here ft: : narrow senseheritabilitl'. or" : additive genetic variance'

o; : digenic geneticvariance.o; : environmental variancedue to replications. and ol : pooled error including environmental varlance
a m o n g p l a n t s w i t h i n r e p l i c a t i o n .S t a n d a r d e r r o r o f t h e h e r i t a b i l i t y
e s t i m a t ew a s c a l c u l a t e da c c o r d i n g t o H a l l a u e r a n d M i r a n d a ( 1 9 8 8 ) .
Statisticalanalysiswas performed using the StatisticalAnalysis S.vstem
( S A S 1 9 8 8 )G L M p r o c e d u r e .
Responseto selection:Three selection methods. differing in parental
'BIC-7-WH' germcontrol and selectionpressure.were applied on the
plasm to determinethe most efficientselectionmethod to increaseseed
S1ZC.

The lirst selection method allowed control over both the selected
seedand pollen parents (full control, FC) and utilized a 10o/oselection
pressurefor both large and small seed.One-hundredand fifty randomly
'BIC-7-WH' were grown and randomly inters e l e c t e dp l a n t s f r o m
c r o s s e du n d e r g r o u ' t h - r o o m c o n d i t i o n s . O n e - h u n d r e da n d t w e n t y o f
t h e o r i g i n a l p l a n t s p r o d u c e d a s u l i l c i e n tn u m b e r o f s e e d ( > 1 5 0 ) f o r
subsequentrneasurementsthrough the DIA for seedsize.Twelve plants
rvith the highest itverage seed size were identified as the large-seeded
p o p u l a t i o n t L S t a n d 1 l p l a n t s w i t h t h e l o w e s t a v e r a g es e e ds i z ea s t h e
s m a l l - s e e d e dp o p u l a t i o n ( S S ) . T h e s e p l a n t s w e r e p l a c e d b a c k i n t h e
g r o n t h - r o o m a n d r a n d o m l y 'i n t e r c r o s s e d( w i t h i n r e s p e c t i v eL S o r S S
p o p u l a t i o n s) $ ' i t h o u t e m a s c u l a t i o n .
T h e s e c o n ds e l e c t i o nm e t h o d a l l o w e d c o n t r o l o v e r o n l y t h e s e l e c t e d
s e e dp a r e n r s( h a l f c o n t r o l . H C ) u i t h a 1 0 7 0s e l e c t i o np r e s s u r ef o r b o t h
l a r g e a n d s m a l l s e e d .T h e m a t u r e s e e dp r o d u c e d o n t h e 1 2 L S a n d 1 2
S Sp l a n t sl i o m t h e i n r t i a l 1 5 0p l a n t i n t e r c r o s sw e r eu s e df o r t h i s s e l e c t i o n

Table l: ANOVA tablefor a genetic

study of alfalfa seed size using
North Carolina Design II mating
d e s i g n( r a n d o m m o d e l ) a n d ' B I C - 7 WH' used as referencepopulatton

Source
Set
Rep (set)
Female (set)
Male (set)
Female x male (set)
Pooled error

df

Fisher's protected least significantdifferencetest.

Results
QuantitatiYe analysis
Seedsizemeasuredon the parental plants involved in the crossing design was not correlated to the seedsize measuredon the
progenyplants (r : 0.256flS,Il : 44). Seedsizeon the parental
plants was influenced only by the seed(female) parent (Table
l). Crossingdateswere used as replicationsand they also had
an influence on seed size. Seed size on the pro-sen)'plants (a
single plant used as the pollen source) was influenced significantly by both the seedand pollen (male) parent involved in
the crossingdesign,and also the interaction betueen the two
(Table 1). Replications in this case were representedu'ith five
full-sibs and did not have a significant effect.
Componentsof geneticvarianceestimatedon parental plants
had values of oi :0.1292 and oi : 0'0084 and showed that
only o2ahad a significant role in determining seed size, which
would be expected with a maternally inherited trait. Despite
t h i s , n a r r o w - s e n s eh e r i t a b i l i t yw a s o n l y 2 3 . 6 9 0 ( S E : 2 2 . 1 ) .
When seed size was measured on the progenr. plants, both
additive and non-additive components of genetic variance
appeared to be involved in the inheritance of seed size
(o2o:0.049 and o2p: 0.049, respectivel))and narrow-sense
h e r i t a b i l i t vw a s 4 1. 3 % ( S E : 1 5 . 1 ) .

Parental plants
MS

2
t2
q
o

21
r61

5.57n
1s
0 . 3 8x5*x
I .416**r<
0 . 1 0 1n s
0 . 0 6 5n s
0 . 0 58

k i g n i l i c a n ta t p : 0 . 0 5 .0 . 0 1 a n d 0 . 0 0 1
* , r ( > kr ,( r . ,S

dI

Progenl plants
MS

2
l2
9
9

:-'\
116

0 . 2 7 3n s
0.026ns
0.416***
0.223*
0.075*8
0.034

EMS
fmo;

^ ' - ) oi..
- +

rmoi
roi- + rfoi

o: -t
6'

r39

A n a l v s i sa n d r e s p o n s et o s c l e c t i o no f a l fa l f a s c c ds i z c

T a b l c 2 : A v e r a g e a l f a l t a s e e ds i z e m e a s n r c da s t h e a r e a o f t l - r es e e d Discussion
i m a g e ( m r n r l a n d s e e dw e i g h t ( g r l 0 0 s e e d s )f o r s c l e c t e da n d u n s e l e c t e i l
P e d e r s e na n d B a r n e s( 1 9 7 3 )p o s t u l a t e dt h a t s i n c ea l f a l f a s e e d
p o p u l a t i o n so 1 ' B I C I - 7 - W H "( R C B D . n i n e r e p l i c a t i o r - r s )

Population

Mean

FC-LS]
FC-SS
HC-LS
HC-SS
NC-LS
NC-SS
BIC-7-WH

3 . 0 2 9a r
2 . 1 1 0c
2 . 8 2 2b c
2 . 6 9 8c d
1 . 8 9 7a b
2 . 5 2 6d
l.7ll5 bc

Size
Gairr

0.214
0.075
0.037
- 0.0u7
0.112
- 0.259
0.261
bc

Wcight
Meern
0 . 2 cI) a
0 . 2 5 9c
0 . 2 6 8b c
0 . 2 5 0c d
0 . 2 8 1a b
0 . 2 3 7d

r F C - L S ( f r - r lcl o n t r o l l a r g e s e e d ) .F C - S S ( f L r l l
c o n t r o l s r r r a l ls c c c l ) t. J C L S ( h a l f c o n t r o l l a r g es e e d ) .H C - S S ( h a l f c o n t r o l s m a l l s e c d ) .N C - L S
( n o c o n t r o l l a r g es e e d )N
. C - S S( n o c o n t r o ls m a l ls c c d ) a. n d ' B I C - 7 W H '
( u n s c l e c t e cplo p u l a t i o n )
r Population means fbllowed by the samc lcttcr
are not significantly
d i l l ' e r e n ta c c o r d i n gt o F i s h e r ' sp r o t e c t e cLJS D t e s t a t P - 0 . 0 5 .

5s11--pollination
did not have a consistenteffecton thc sizc
of the resultingS, seed.Measurernents
of seedsizeafter crossand self-pollinationon 48 individual plants showedthat self-ed
( S ' ) s e e dw a s s m a l l e rt h a n F , s e e di n c e r t a i nc r o s s e sb. u t a l s i r
l a r g e r t h n n F , s e e di n o t h e r c r o s s e sT. h e r e w a s a s i g n i f i c a n t
c o r r e l a t i o n( r : 0 . 1 2 1 ' " * * : P < 0 . 0 0 1 :n : 4 6 ) b e t w e e nt h e s i z e
o f S , a r - r dF , s e e dd e v e l o p e do n i n d i v i d u a lp l a n t s .f u r t h e r c o n Iinning the importancc-of the seedparent in detel.rining seecl
SIZC.

Responseto selection
B a s e do n i n d i v i d u a l p l a n t r l e a n s . t h e i n i t i a l b a s ep o p u l a t i o n
( 1 2 0 p l a n t s .' B I C - 7 - W H ' ) f o r s e l c c t i o nm e t h o d so n e a n d t w o
(FC and HC) had an lrvera-seseed size of 1.633nrmr and it
s t a n d u r dd e v i a t i o n o f 0 . 2 3 6 . w i t h a s l i g h t s k e w t o w a r d t h e
h i - c h e rv a l u e s .T h e o r i g i n a l 1 2 s e l e c t e dp l a n t s .b o t h f b r l a r g e
ar-rdsmall seed.had an averageseed size of 3.058rnmr and
2 . 2 9 6r n m r .o r s e l e c t i o d
n r f f e r e n t i a l o0f . 4 2 5m m r a n d 0 . 3 3 7 r n m t .
respectivell,.
A t l e r o n e c l c l e o f s e l e c t i o n t. h e t h r e e s e l e c t i o nm e t h o d s
resulteclin ditl-erentselectiongains (Table 2). There wlls a sign i f i c a n ts c p a r a t i o nb e t u e e np o p u l a t i o n ss e l e c t e d
f o r l a r g ea n d
s m a l l s e e d w i t h t u o o f t h e s e l e c t i o nm e t h o d s ( F C l a n d N C
populations). There u as also a significant seed size shift
b e t u e e n t h e u n s e l e c t c dp o p r - r l a t i o(n' B I C - 7 - W H ' ) a n d F C I - L S
and NC-SS. confirming tliat alfalla secd size was Lr heritable
trait fbr the relerencepopulation.
The differencesfor realized gairr from different selection
n-rethods
suggestedthat parentalcontrol was intportant for the
successof seeclsizeselectionin allalfn. especiallyfor increased
s e e ds i z e .T h e o r e t i c a l l yH
. C - L S s h o u l d h a v e r e a l i z e da p p r o x i rnatelyhalf as much gain for sccdsizeas FC-LS, sinceselection
for these populzrtionsdiffered onlv in parental control. The
pressllre
i n t e n s es e l e c t i o n
o f < l % r a p p l i e dl o r t h e N C - L S p o p L t lation did not compensatelbr lack of parentalcontrol for lar-ee
gain Ibr smaller seecl
seed.but NC-SS did sliow'cor-rsiclerable
size.
Seed w'eight \\1us strongll' associated with seed slze
( 1 : 0 . 9 7 8 * * * ; P < 0 . 0 1 : n : 7 ) . a n d t h e r e l b r et h e s e l e c t e d
populationsshowedthe santedifl-erences
for seedlveightas they
did for seedsize(Table 2).

had essentiallyno endosperm.seed size should reflect or-rly

embryo sizeand therefbrecould serveas an indication of seed
hybridity. They reportedthat hybrid seed(Fr) wele : 5% larger
than sibbedseed,and sibbedseedwere :5'% largerthan selfed
( S , ) s e e d .B o w l e l ,( 1 9 8 0 )f o u n d t h a t s e e dw e i g h tw a s g r e a t e ri n
S, than in F, seed.He suggested
that reducedf'ertilizationcaused
by self-incompatibilitymay have reducedthe number of S, seed
a n d r e s u l t e di n l e s sw i t h i n - p o dc o r n p e t i t i o n .
Morc rccently. a model of geneticcontrol o1'seed weight
was established
b y P e t e r s o na n d B a r n e s( 1 9 8 2 ) .T h e i r r e s u l t s
suggested
that seedweight was controlledprimarily by additive
gene actior.r.and scedlingvigour was controlled prin-rarilyby
nor-r-additivegene action. They also reported a significant
maternal (seedparent) effecton seedweight. Ciornplernentary
r e s e a r c h( K a t e p a - M u p o n d w ae t a l . 1 9 9 6 )a l s o s h o w e dt h a t t h e
influenceof the seedparent on sccdweight wils morc consistcnt
t h a n t h e i n l l u e n c co f t h e p o l l e n p a r e n t .
Resultsof this reseurchindicatetl"ratboth additiveanclnonadditivegeneacticlndeterminealfalfa scedsizc.and also suggcst
t h a t i t i s n o t a p p r o p r i a t et o r n e a s u r et h e - q e n e t iecx p r e s s i o n
for
seedsrzeon seeddevelopedon the parentalplants involved in
a crossingdesign.Seedsize should be measuredon seedproduced on the progeny plants. In other words. seedsizeis cot.rtrolleclby maternal plant genotypeto a greaterextent than bv
the genotypeof thc embryo comprisingthat seed.
Sclcction for both large ancl small seed resulted in a signilicant responseafier only one cycleof selection.but the selection was not equallyeffectivcrn both directions.Asymmetrical
responsescornmonly occur in the first cycle of bi-directional
s e l e c t i o n( F a l c o n e rl 9 8 l ) . T h e a p p a r e n td i f f e r c n c ersn s c l e c t i o n
gain for largeand srnallseedwithin the sameselectionrlethod
can be attributed to severalfactors.including:non-nonnal distribution of tl"reinitial population. andior difi'erentinheritance
mechanismsassociatedwith the largeand srnallscedtrait. Heterozygosity.or the presenceof tri- and tetra-allelicloci. lias
been reported to marimize seed size (Duribicr ar-rdBingham
1915).Therefbre.it rvor.rld
bc very unlikely that heterozygosity.
usuallyassociatedwith vigour. could act toward reducingseed
sizeA
. p l a u s i b l eh y p o t h e s i iss t h a t o n l y a d d i t i v eg e n e t i cv a n a n c e
d e t e r m i n e s m a l ls e e da n d t h a t b o t h a d d i t i v ea n d n o n - a d d i t i v e
gerieticvariance(heterozygositl,)
determinelarge seed.
In conclr,rsior-r.
this researchindicatedthat for geneticstudies.
seedsizeshould be measuredon the seedproducedon progenv
plants.For the rcfercnccpopulzrtion.allalfa seedsizewersshown
to be a highly l"reritable
trait. Selectionfor increasedseedsize
rvaseffectiveaftcr one cvcleof selection.

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