Jf nstttutt for
grønsak~yrking
Stensiltrykk nr. 156
GENETIC VARiATION AND INHERITANCE OF CERTAIN
QUALI TY CHARACTER I ST I C S AND I TS U SE
IN BREEDING OF CABBAGE
By
Magnor Hansen
Forelcsninq ved niende fel!esnordisl<e lic/doktorandkurs i
planteforedling 17-22 januar 1983, Dan mark
GENETIC VARIATION AND INHERITANCE OF CERTAIN QUALITY CHARACTERISTICS AND ITS USE IN BREEDING OF CABBAGE
By
Magnor Hansen
Forelesning ved niende fellesnordiske lic/doktorandkurs i
planteforedii ng 17-22 januar 1983, Dan mark
- 2 -
GENETl.C VAR.lATION AND I.NHERITANCE OF CERTAlN QUALITY CHAMCTEIUSTICS AND ITS USE IN BREEDING OF CA,BBAGE Magnor Hansen
Department of Vegetable Crops·
Agricultural University of Norway P.O.Box 22, N-1432,Aas-NLH
Norway
In future breeding plans of Norwegian c~bbage, we have set our sights on a number of goals (NLVF-report no. 109). Three important characters concerned with quality are:
1. - Storage qualities
2. Length of the stem inside the head (Inner stem) 3. Length of the stem above the ground (Outer stem) Storage quality isa very important quality character because the aim, in the future, is that Norway should be self-sufficient with cabbage for most part of the year.
With regard to the length of the inner stem, it is desirable that this is short. A lang inner stem gives rise toa less compact head and thus a lower quality.
It is also probable that future breeding programmes will include demands on the size of the outer stem. If the outer stem is too lang, cabbage plants have e tendency to lodge and therefore making harvesting more difficult. On the other hand, if the outer-stem is too short, this would result in the head touching the ground and making it susceptible to so~l pests and diseases.
It is also possible that future mechanical harvesters will set special demands on the length of the outer-stem.
Materials and methods
To study the interitance of a number.of characters in cabbage, The Department of Vegetable Crops, at the Agricultural University of Norway laid out a 10 x 10 crossing where 10 genotypes were crossed diallelic {Table 1).
Genotype 0-7 originate from the Department of Vegetable Crops own breeding material. Genotype 8 comes from the American variety 'Badger Shipper',while the last genotype is from the Norwegian commercial variety 'Aglo'.
Befare starting on the diallel crossings, all the genotypes were cloned from leaf shoots (NORTH 1952, SNEDDON 1962, HANSEN 1981).
None of the parental clones, except clone 8, were inbred befare they were included in the present experiment. The
diallel cross was complete, that mearis that all the individual combinations were fertile.
In 1980, plants were produced from each crossing. They were planted out in a randomised block experiment. In addition to the 100 F
1 families, the 10 parental clones were also vege- tatively propagated and included in the block experiment.
Each plot had two rows o~ 6 plants spaced at 50 cm between plants and the rows were set 60 cm apart.
A number of observations were made on several different charac- ters. However, from here on we shall be concentrating on the three characters that were rnentioned in the introduction.
The sto~age quality was estimated by storing 4 heads from each
-. 4 -
of the F
1 families and parental clones in a cooling rom. The heads were weighed befare storage. During the storage period the heads were taken out, cleaned and weighed and put back again. This was done after about 75 and about 150 days of storage. Results show that the saleable yield was reduced
linearly with storage time (Figure 1) ~ As a measure of storage quality, the percen~age of fall in the saleable yield after 100 days was calculated (Figure 1). The storage experiment has 4 replications.
The length of the inner stem was rneasured at harvest. 8 heads from each
r
1 family and parental clones were divided verti- cally and the inner stem rneasured. This experiment had 5replications.
The length of the outer stem was measu~ed on all the plants in the block experiment two weeks befare harvest. This experi- ment had 5 replications toa.·
Results
In the figures 2, 3 and 4, the distribution of family means for storage quality, length of the inner stem and length of the outer stem are presented in a graphic form. The diagrams showa good approach toa nearly normal continuous distribution.
This continuous distribution shows that these three characters must be considered as quantitative characters. In other words,
there must be a number of genes that control these characteres.
The regression of covariances between the diagonal families and arr~ys (W
21 )
on variances of arrays (V) is deduced byp r r
DICtINSON & JINKS (1956}. Figure 5 shows the Wp
2/r/Vr - graph for storage quality. From this graph we can see that the
genotypes 5, 6 and 9 have most dominant alleles with regard to storage quality. In addition, the regression line cuts the w p2/r - axis above origo. This means that the char.acter for
storage quality shows incomplete dominance.
The W
21 .IV - graph for the character length of the inner
p _r r ·
stem (Figure 6), shows that also this character shows inqomplete dominance. Genotype 9 has most dominant alleles for this
character.
From the W
21 /V - graph for the length of the outer stffin
p r r
(Figure 7), one can see that this character too shows incom- plete dominance, and that the genotypes 2 and 9 have most dominant alleles.
The W
21 /V - graphs show that genotype 8 is the most recessive p
r
rfor all the three characters.
Table 2 shows the expectations for array variance (V}, cova- r
riance between parent clones and arrays (Wpl/r) and covariance between inbred. lines (diagonal families) and arrays (Wp
2/r), in a diploid heterozygous diallel cross.
I have found a linear regression between Wpl/r and wp 2/r·
expectation for the regression coefficient is:
The
bwpl/r/wp2/r
= =
wpl/rAa wp2/rAa
=
Wpl/raa w/ p2 raa
d + h(l-2u) - d + \h(l-2u)
The slepe of the regression line is determined by the level of dominance and the gene frequency. From the regression coefficient one can see that when the gene frequencies are equal (u = v = 0.5), the regression coefficient is equal to 1.
We can also see that when u is greater than
v,
the regression coefficient becomes less than 1. The regression coefficient~
is lar.ger than 1 if u is less than v.
Figure 8 shows the W l/
_/W 21 _ _ -,
graph when h = ~d for thep r p r ·
gene frequencies u
=
0.2, 0.5 and 0.8 respectively. Figure 9 shows the Wpl/r/wp2/r -, graph when h=
d for the same gene frequencies, while figure 10 shows the Wpl/r/Wp2/r - graph when h
=
d and gene frequencies u=
0.2, 0.5 and 0.8 respec- tively"We see that irrespective of the level of dominance or the gene frequency, we can expect the regression line to go through the point of origo. At full dominance (h = d), the observations are expected to circle around origo. In the case of over-·
dorninance (h > d) ,_ the observations are expected to fall in the third quadrant, while at incomplete dominance (h < d), the observations will arrive at a point above origo. Observations for the most recessive genotypes will in all cases be located furthest away from origo in the first or second quadrant.
With incomplete dominance, an alteration in the gene frequency will lead to only a small change in the ~gression coefficient.
If his large, an alteration in the gene frequency would lead toa larger change in the regression coefficient. If h
=
0,there will be no significant regression at all.
Figure 11 shows the Wpl/r/wp2/r - graph for stor.age quality.
The distribution of the points along the regression line shows incompJ_ete dominance. This graph also shows that the genotype 5 has ~ost dominant alleles while genotype 8 has most recessive alleles. The Wpl/r/wp
2/r.,.. graph for the character length of
the inner stem (Figure 12] shows that genotype 9 has most Aominant alleles for this character. It is also genotype 9
that is most dominant with regard to the character length of the outer stem (Figure 13). Geriotype 8 is most recessive in all of these three characters. The regression coefficients show that there isa surplus of dominant alleles for all the three characters.
In tables 3 and 4, DR, HR and the heretability in broad and narrow sense are estimated from the variance components
(AASTVEIT 1966, MATHER & JINKS 1977).
The DR's is estimated from the variances of array means (Vr), the mean covariances between par-en t a I clones and arrays (Wpl/r) and the covariances, between the rneans of parental clones and the means of .their offspring fam~lies {Wpl/o). The HR's are estimated by substitution in the equations for the variances of parental clones (Vpl}, the mean variances arrays (Vr) and the variances of offspring families around the total means
(V-) •
0
Conclusions
The results from the analysis of the diallel cross show that the three characters in question are inherited quantitatively, and that a large part of the variation is additive. One can thus expect a great response to selection.
It should be possible, from the progeny of the diallel cross, to create new varieties with storage qualities that are above average. Following this, one can carry out stabilized selec- tion for the characters for length of the inner stem and length
- 8 ...,
of the outer stem, so th.at these characters ~re expressed properly in new varieties.
As can be seen from the results, the genotype 8 (from 'Badger Shipper') is the most recessive for all the three characters.
However, genotype 8 is evidently the most recessive genotype for all the characters examined in this diallel experiment.
From this, we can expect that this genotype should be a suit- able cornmon tester if one wanted to use the top cross as a method for progeny testing.
References
Aastveit, K.' 1966: The Value of Biometrical models in Plant Breeding.
· Acta Agricultura Scandinavica, Suppl. 16 (1966)
Chiang, B.Y. & W.F. Grant, 1975: A putative heterozygous interchange in cabbage (Brassica oleracea VAR. capitata) cultivar 1Badger Shipper'.
Euphytica 24: 581-84.
Dickinson, A.G. & J.L. Jinks, 1956: A generalised analysis og diallel crosses.
Genetics 41: 65-78.
Hansen, M~, 1981: Genetic variation and inheritance of tolerance to clubroot (Plasmodiophora brassicae Wor.) in cabbage.
Proceedings of Brassica conference 1981, 15-18th September 1981. Department of Vegetable Crops, Box 22, 'N-1432 As-NLH, Norway. Stensiltrykk nr. 153: 135-142.
Mather, K. & J.L. Jinks, 1971: Biornetrical Genetics.
London, Chaprnan and Hal 1 Lt.d ,
- 9 ,....
North, C., 1952: Vegetable propagation of c~bbage and allied vegetables.
Emp. J. exp. Agric. 20: 43-47.
Snedden, J.L., 1962: A note on vegetative propagation of some forms of Brassica olerace~.
J. nat. Inst. agric. Bot. 9: 145-48.
Weisæth, G., 1948: Utvikling av klumprotresistente kålsorter ved kombinasjonsforedling og gjentatt seleksjon på
Plasmodiophora-infisert jord.
Forskn. Fors. Landbr. 19: 233-54.
- 10 -
Table 1. Parental clones in the diallel cross.
Geno-
type Origi n References
0 1 2 3 4 5 6
·7 8 9
'Reslsta' ('Jåtunsalgets v. k. ' x 'Bøhmer- wald')
li
li
li
li
TK 704 I'Rosseba' x 'Bøhmerwalo') x
1
Bi ndsachsen'
li
K 707 ('Rossebø' x
IBøh m_erwald') 'Badger Shipper' (Brassica oleracea capitata x B. o. acephala)
'Agio' (Selected in 'Toten Amager')
Weisæth 1968
li
li
li
li
li
li
li
Chiang & Grant 1975
- 11 ~
YIELD, KG
- .-1 % LOSS IN WEIGHT AFTER 100 DAYS
IS
DEFINED AS STORAGE QUALITY
20 60 100 140 180
STORAGE TIME
(DAvs)
FIG. l
IESTIMATICN OF STORAGE QUALITY
No, OF PLOTS
90
. 70- 50
30
10
•
i"'
~
P"
. .
~
t"
~
..
~
..
STORAGE QUALITIE
. . •. s
5 · 15
10 20 25 _30 35 45
40 50
FIG.
L ·•· D
1 STRr
BUT 1 ON OF STORAGE QUAL 1·r1 ES55 60 65
-. 13 -
No. OF PLors
90
70 ·
50
30
· 10 -
I
I . I I
,
,,..-- ..
8')5 10.25 11')5. 13·.·25
I J- - • - - - - I -- - •••.•• I l14,75 16,25 0
LENGTH OF THE INNER STEM (CM)
FIG·,·
3 •·
DI STR .I.BUT ION
OF LENGTH OF THE STEM I NS I DE THE· HEADNo. OF PLOTS
90
- 70 50
30
10
'.
"'
...
..
..
..
.
"'
.
..
..
""
I I
I I I
I I
..
3 5 7
4 6 9 11 13 15 _
8 _ 10 12 14 16
LENGTH OF THE STEM ABOVE GROUND (CM)
fIG. 4 , IlISTRIBUTION OF LENGTH OF THE STEM ABOVE GRCUND
- 15 -
200
100
/ 8
WP 21 r
=61, 6 7 + 1, 02 V r
R ~ 0, 9 581 b
=1. 02 ± 0. 1
100 200
Fig. 5. (Wp2/r/V r) - graph for storage quality
300
200
100
9 •
wp 2 /r
=61, 44 + o, 924 v r
R
=0, 9283 b
=0. 924 ± 0. 13
,, 8
100 200 300
Fig. 6. (Wp2/r/Vr) - graph for length of the inner stem
~, 17 ~ .
• 3
2
1
•O
wp 2 /r
=o, 994 + 1, 008 Vr
R
=0, 9 5 28 b
=L 008 ± 0. 11
--
N' I-~= I
I1 2 3
flg. 7. (Wp2/r/V r) - graph for length of the outer stem
-. 18 -,
Table 2 • Variances of arrays (V r), covariances between parents and arrays (Wpl/r) and covarlances between the diagonal and arrays (Wr /r) in a heterozygous diallel cross.
2 '
Genotype
Fenotype vr Wpl/r Wp2/r
Frequency
AA ½UV(d-h) 2 uv(ct2-2udh uv(ct2- ½ (1+2u)dh
d +(l-2v)h 2) - + ½.U-2v)h2)
u2
AA ½uvd2 uv(d2 + (v- u)
Iuv(ct2+ ½(v-u)dh)
h dh)
2uv
aa ½UV(d+h) 2 uv(ct2+2vdh uv(ct2+ ½(I+ 2v)dh
-d +(l-2u)h2)
T½ (l-2U)h2)
v2
wpl/r AA wol/r Aa wpl/r aa d+h(l-2u) bwpl/r/Wp2/r - Wp2/r AA = w = w
=p2/r Aa p2/r aa d+ ½h(l-2u)
\
- 19 -
0,4
--
L- r--Cs:
0..0, 2
u=O, 5,
b=l
-,
u=O 2
Ib=l, 1297
aa
u=O 8 , b=0,82?5
0, 2 Wp2/r
0,4
Fig. 8. The relation between W l/ and W
21
in ap r p r
one-gene heterozygous diploid diallel cross.
h = ½d. u = 0.2, 0.5 and 0.8.
- 20 -
0 4
i-l
0, 2
u=-0 2 ,
b=l 3 , u=O 5 b=l '
aa
Wp2/r 0,2 0,4
Fig. 9. The relation between Wpl/r and wp 2/r heterozygous diploid diallel cross.·
h
=
d. u=
0.2,o.s
ando.a.
in a one-gene
- 21 ...
s....
-.. _.
CL.
$
0,8
0,6 0,4
0,2
-0 2 ,
AA AA
u=O , b=l,375
aa u=O 5 , b=l
0,4
u= O 8 ,
b=
-.Q, 5
aa
0,6 Wp2/r
Fig. 10. The relation between W l/ and W 2.
1
in a one-genep r p r ·
heterozygous diploid diallel cross.
h = 2d. u
=
0.2,o.s
and 0.8.- 22 •....
200
100 ~
-- ,._
,---.1
c..
$
WP 11 r
=3, 47 + 0, 6 2 WP 21 r
R
=0, 9672 b=O. 62 + 0. 06
wp2/r 100 200
Fig. 11. (Wpl/r,wp2/r) - graph for storage quality
-. 23 't",
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