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Molecular markers for cytoplasm
Male sterility and contribution of different plastid-mitochondrial
configurations to starch production
A. Lössl 1, 2, M. Götz 1,
A. Braun 1, G. Wenzel 1
1 Lehrstuhl für Pflanzenbau und Pflanzenzüchtung, TU
München-Weihenstephan, D-85354 Freising (Germany),
2 Institute of Botany, Ludwig Maximilian University, Menzinger
Str.67, München (Germany).
Distinct parental cytoplasms were combined in symmetric tetraploid
hybrids of potato by somatic cell fusion. This allowed, in the presence
of nearly isogenic nuclear genomes, to estimate the contribution of mitochondrial
(mt) and chloroplast (cp) genomes to starch content. Analysis of mt-cp
configurations in the complete gene pool of german potato cultivars [2n=4x],
in a reciprocal dihaploid population [2n=2x], in di-haploid fusion parents
[2n=2x] and in their respective hybrids [2n=4x] made visible the effects
of different cytoplasmic backgrounds and mitochondrial subgenomic rearrangements.
Genotypes identified by markers as cytoplasm Wg
were associated with cytoplasmic male sterility. Evaluation of cytoplasmic
types lead to the conclusion, that in starch content the ´wild type´
Wa and Wg
have a significant advantage to other cytoplasmic types (T
b,W d and S e).
This results from the experiments with a reciprocal population, 180 di-haploids,
and from cultivar comparisons.
In hybrids an interaction between starch content and different mt-cp
combinations could be found.
In general the highest field performance, measured in starch content
and yield was associated with such cytoplasmic configurations which appeared
to a high frequency within a population, when the segregation process was
completed. This fact is explained by a selection advantage of clones
with optimized organellar segregation already during in vitro phase. PCR
markers for cytoplasm differentiation are actualized on a website.
Key words: potato, plastid, cytoplasm, male sterility, starch,
In most cases the influence of the type of cytoplasm is not
distinguished from biparental inherited traits. Moreover, cytoplasmic
phenotypes only occur in combination with distinct nuclear back-grounds
and this hampers their recognition in further breeding work. Unintended
counterselection against some organellar types may take place, as long
as the agronomic value of different cyto-plasms remains unknown.
Information about these influences is valuable for selection in sexual
breeding but also for production of somatic fusion hybrids.
Following determination of different cytoplasmic types, cytoplasmic,
and in part maternal effects can be detected in at least three ways:
1.) Comparison of gene pools of clones (4X cultivars), di-haploids
(2X breeding clones),
In conventionally bred clones an optimized nuclear-cytoplasmic interaction
can be achieved in several traits. This was proven by experiences of conventional
plant breeding where reciprocal crosses show differences between cytoplasms:
in this respect differences in the photoperiod reaction of different plasms
(Sanford and Hannemann 1982) are relevant as well as maternal effects,
reported by Maris (1989) who analyzed a series of diallel crosses between
ssp. tuberosum and adapted ssp. andigena varieties: populations with tuberosum
cytoplasm were found to be superior in respect to tuber yield whereas the
andigena cytoplasm provided a higher male fertility. Hilali et al. (1987)
observed reciprocal differences in tuber yield, tuber number, vine vigor,
average tuber weight, seed germination and pollen vitality. In cytoplasmic
substitution backcrosses of potato Amoah et al. (1988) found reciprocal
differences between S. phureja and S. tuberosum backcross
2.) Comparison of reciprocal populations, and
3.) Comparison of somatic hybrids containing different cp/mt configurations
with nearly identical nuclear genomes.
Nevertheless maternal effects on field performance which are dependent
on interorganellar energy metabolism as is starch accumulation and degradation
have not been reported so far. This might have been due to the lack of
markers capable to differ between cytoplasmic types.
In conventional breeding the differences in sterility, tuber characteristics,
seed production and germination are conditioned unlikely by cytoplasmic
factors alone. For this reason it is useful to analyze symmetric hybrids,
which possess isogenic nuclear genomes and different cytoplasmic backgrounds.
By this constellation the effects of various cytoplasms and of plastid-mitochondrial
(cp-mt) interaction can be separated from nuclear differences.
In contrast to sexual combinations in somatic fusion the genetic information
of the cytoplasms is biparentally inherited. Therefore it is relevant to
investigate the influence of the different cp / mt configurations after
segregation within a hybrid population, generated in fusion experiments
(Schilde-Rentschler et al. 1995, Cardi et al. 1999).
Field trials show variability between hybrid clones from the same parents.
This variability can be due to nuclear deviations and different cytoplasmic
configurations. The latter were analyzed in detail in an investigation
of the fate of organelles after cell fusion by Lössl et al.
(1999). The characterization of potato cytoplasms in five main cp/mt
types Wa T
and S e was a precondition for the detection
of correlations between cytoplasm and phenotypic data. A set of probes
and PCR primers useful for the quick characterization of breeding material
was developed and actualized on a website.
Novel configurations of cytoplasmic types occurred in fusion hybrids
in shape of partial or complete additions of parental fragments and even
novel fragments. In fusion hybrids the strict assortment of cp and mt types
was lacking; within regenerants of potato, plastids segregate completely
into one of the parental types, whereas mitochondrial genomes are affected
from various portions of rearrangements. Cp/mt differentiation serves as
a basis for a subsequent evaluation of cytoplasmically influenced traits.
In symmetric hybrids with isogenic nuclear genomes - neglecting protoclonal
variation - it could be shown that in general a low degree of chondriome
recombination correlated positively with yield potential (Lössl et
al. 1994). The question arose, whether mitochondrial performance is influenced
by distinct chondriome regions and whether they can interact more efficiently
with the new generated genome if they derive from a special parent (Lössl
1996; Frei et al. 1998). An evaluation of traits being influenced by cytoplasm,
is expected to allow predictions about the effects of distinct cytoplasmic
configurations for breeding purposes.
Material and Methods
Restriction fragments specific for the different cytoplasmic types
(a, b,g,d,e, k)
were cloned and sequenced according to Sanger (1977). The potato mtDNA
probes, used in this work can be supplied on request. PCR primers specific
for mitochondrial types (a, b, g)
were designed, with the pairs of primers used to detect mt type g
associated with cytoplasmic male sterility (CMS). They are given in Table
Nearly the complete spectrum of the german cultivar sortiment and 180
dihaploid clones were analyzed on their cytoplasms. Using 4 pairs of PCR
primers, and 11 homologous mt-probes they were grouped as mt types a,b,g,d,e
Optimal annealing temperature for mt primers was 57°C, and for
cp primers ALC_1 and ALC_3, to differ between cp type T and wild type plastids
(cp type W or S) 44°C. The markers were used for the estimation of
cytoplasm frequencies within the german potato gene pool (Table 2). Plastid
type nomenclature was used according to Hosaka and Hannemann (1988).
The fusion hybrids have been checked for their deviations in their
nuclear genomes by usage of a set of probes, which covered the 12 chromosomes
as performed by Lössl et al. (1994).
For the present analysis fusion populations which yielded about 30
hybrid clones, each going back to a single fusion event, were chosen. Fusion
hybrids and the dihaploid material derived from the Technical University
Munich and from the Institute for Resistance Genetics (Bundesanstalt für
Züchtungsforschung) in D-85461 Grünbach. A reciprocal F1 population
(Table 3) was provided by Dr. Uhrig, MPI Cologne. Fusion
populations employed in this work are listed in Table 4.
Field experiments, performed during four years, consisted of blocks
in two and three replications with approximately 3,5 m2 and 16 plants per
field plot. Data were evaluated at two locations: in the experimental fields
of the Institute for Resistance Genetics, Grünbach and at Roggenstein
(Oberbayern) and of the Technical University Munich. Populations (Pop),
which were basis for highly significant observations (P=0,01), consisted
of more than 30 hybrids. Pop Ia [FAL2 (+) 601] consisted of 35 symmetric
hybrids, grown in nine replications, Pop Ib [BP32 (+) 601] of 55 hybrids
in two replications, Pop Ic [2006 (+) 626] of 6 hybrids in five replications,
Pop IVb [FAL2 (+) M9] of 32 hybrids in two replications, Pop IIc [26-2
(+) 1512_25] of 17 hybrids in five replications, Pop IIIb [2006-10 (+)
576-16] of 10 hybrids in five replications, Pop IVa [BP32 (+) M9] of 58
hybrids in 6 replications, where two plots were lost. They were calculated
as an imbalanced trial. Additional field data for the various cultivars
were given by the German Seed Board from the “BundesSortenamt”. Starch
and dry matter was determined using a starch weighing machine type MEKU
(Pollähne, D-30974 Wennigsen, Germany). For the calculation of the
influence of different cytoplasmic compositions on yield components ANOVA
and non parametric statistical ´SPSS´ programs for imbalanced
groups were used.
Test for pollen fertility and vitality was carried out by staining
procedures and germinating tests respectively (Stanley and Linskens 1974).
Differentiation of Cytoplasms
Within 4X cultivars predominantly the cp/mt types Wa
occurred. Table 2 gives the percentages for this mt
type distribution within the German cultivars.
By usage of DNA markers (probes and PCR primers) it is
now easy to differ between the cytoplasmic types of potato mt-a,
b, g. Figure 1 shows a comparison
of cytoplasmic types Wa T
with the specific PCR
primers. The primers ALM_1, ALM_3, ALM_4, ALM_5, ALM_6 and ALM_7 in combination
with ALC_1 and ALC_3 as well as the probes m79, m80, m93, and m112 can
be applied as markers. They detect mt type g,
which is associated with the phenotype of cytoplasmic male sterility. The
set of probes for determination of mitochondrial and plastid genome
types was tested in non-radioactive hybridization. In this way breeders
can perform DNA analysis without isotopic laboratory.
Comparison of field data to cytoplasmic configurations
In order to detect advantageous effects of distinct cytoplasmic types,
the di-haploid genotypes, hybrid populations, the reciprocal F1 population
(US1) and the whole spectrum of German cultivars, were grouped according
to their organellar types and compared in agronomically interesting traits.
For this purpose their starch contents, yields and fertility were evaluated
(Table 3, Table 4). Cytoplasmic
male sterility (CMS) was confirmed by pollen germination tests. Most of
the CMS genotypes could be traced back to ancient genotypes as are: I-301,
MPI 61-303/34, 56.4129/288, MPI 46.956/68 and Röslau. This maternally
inherited sterility was correlated with clones containing mt type g,
as are Aiko, Alwara, Assia, Azur, Bettina, Forelle, Fox, Heidrun, Helios,
Petra, Sibu, Uno and Ute.
Cultivars and di-haploid breeding clones
Correlation of field data with cytoplasmic types showed differences
between the cytoplasmic groups. Comparison of the cytoplasmic types by
Kruskal-Wallis one-way-anova resulted in significantly higher starch contents
of varieties with Wa and Wg
type than those with Tb types (P=0,01).
The effective difference between culture type (T b)
and wild type (Wa, g) was 14,7% vs. 16,2%.
This clear difference was reduced, but still existent, if starch yield
was considered (P=0,05). With slight deviations the field data, measured
in this trial, were consistent with the data given by the German Seed Board
(“Beschreibende Sortenliste“). In order to exclude a coherence between
cytoplasmic type and nuclear genome we performed a cluster analysis by
usage of nuclear genomic probes and RAPD (data not shown). Genetic distance
of nuclear genomes of 144 varieties did not correlate with their assortment
to different cytoplasmic groups.
According to breeders experience and pollen vitality tests nearly all
cultivars identified as mt-g are male
sterile. The probes and primers listed in Table 3 can
be applied as CMS-markers.
Similarly as found in the cultivar pools, comparison of the cytoplasmic
pools within the di-haploid clones confirmed, that the mt-b
pool had significantly lower starch contents than those with the "wilder"
plasms mt- a,g,d,e. In contrast to the
varieties this was associated with a shortened maturity.
In starch content the reciprocal crossed F1 population ´US1´
revealed a significant positive effect of cytoplasm mt-a
mt-d. In view of 41 and 35 individuals
per cytoplasm pool, it was empirically sure that the different nuclear
alleles were represented proportionally within the two groups mt-a
and mt-d. Thus they were comparable.
The effective difference was 14,4% (mt-d)
to 15,8% (mt-a).
For a deeper elucidation of nuclear-cytoplasmic-interactions it was
useful to analyze populations of somatic hybrids, with nearly identical
nuclear genomes, but containing different plastid-mitochondrial configurations.
In hybrids generally the highest field performance was associated with
such cytoplasmic configurations which were represented to a high frequency
within the concerned fusion population (e.g. Ia, Ib, IIc, IIIb, IVa). Clones,
which turned out to have the highest productivity, have already been accumulated
naturally during in vitro phase. This selection pressure was extremely
obvious in population IIIb [W/a (+) W/d]
were 2/3 of the individuals reverted to the configuration of mt type b
at rps14 gene locus.
In contrast to this reconfiguration, hybrids of population IVa revealed
a high correlation between homogeneous mt genomes and a high yield potential
(Lössl et al. 1994). Continuous evaluation of yield data from 1994
to 1998 detected that an irregular amplification of an additional atp6
copy was correlated with severe depressions in starch yield. In population
IVa, these 10% of hybrids, which were affected by a highly replicated additional
atp6 copy (Lössl et al. 1999) had a significantly lower starch content
and yield level than those which did exhibit a normal mt genome organization
Within fusion population Ia plastid segregation ratio was skewed (cpW:
cpT = 1: 6). In starch content hybrids containing cp type T revealed a
clear advantage. But in respect to the novel organization of the
hybrids mt genomes the cp pools had to be subgrouped into mt recombination
types R1 and R2 (Figure 2). In dependence of the mt
type, differences could be detected in starch yield between somatic hybrids.
In connection with cpW the mitochondrial recombination type R1 was superior
to type R2. This correlation was inverted if mt types R1 and R2 were connected
with cp T. The interaction between mt- and cp-types was significant (P=0,001).
This interdependence in starch yield is shown in Figure
The working hypothesis of a correla-tion between cytoplasmic
genome configuration and field performance measured by starch production
could be confirmed in the current investigation.
Evaluation of plasm types lead to the assessment, that in starch
content the ´wild type´ cytoplasms (W/a
and W/g) are superior in comparison to
other cytoplasmic types (T/b,W/d,
This results from field trials with di-haploids, cultivars and a reciprocal
cytoplasm is associated
with male sterility and for breeding of a high starch variety W/a
and W/g appears to be the appropriate
It was generally observed that those cytoplasmic configurations,
which appeared to an unproportionally high ratio in somatic fusion hybrids,
were also associated with the highest field performance. This fact could
be explained by a selection advantage of such clones, which works already
during in vitro segregation and regeneration phase.
Conventionally bred cultivars
Field trials with varieties showed, that the advantage in starch production
of cytoplasms W/a and W/g,
representing the wild type cytoplasms was slightly reduced when starch
yield was focused. Higher yields seem to compensate the lower starch contents
to some extent. The correlation to starch content had a high significance
(P= 0,01), whereas correlation of the same cytoplasms to starch yield was
slightly lower, but still significant (P=0,05). If starch content and yield
are considered as complementary breeding aims, the cytoplasmic advantage
of W/aand W/g
remains relevant for breeding a high starch cultivar.
It is suggested that cultivars with cytoplasm T/b
are optimized for stable yields rather than for high starch contents. Definitely
nuclear genes determine starch content and yield, but during starch accumulation
and degradation they interact with factors coded by the plastid and mitochondrial
genomes. For this reason the complete assortment of cultivars were in parallel
investigated in view to their diversity on nuclear genome level. Following
an association to cytoplasmic pools W/a,
W/g and T/b
the cultivars were tested, whether their cytoplasmic wild type content
corresponds with different portions of wild type character in their nuclear
genomes. So far there is no indication that varieties with the same cytoplasm
could be grouped in the same cluster on nuclear genome level. Nuclear alleles
seem to be distributed rather independent from cytoplasmic mating barriers.
An exception might be the cytoplasm of W/g,
which is correlated with male sterility.
Mt type g has been introduced into
the German spectrum of cultivars together with PVY-virus resistance. For
this purpose the maternal parents from MPI I-301 or its relative MPI-61-303-34
have been employed. These genotypes derive from a mating with a S. stoloniferum
accession, containing a deviating wild type cytoplasm. Somatic fusions
with a parent of mt type g generate hybrids
with different portions of this CMS conferring cytoplasm. Preliminary results
with the somatic hybrid populations showed, that this correlation can be
broken by the novel cp-mt-configurations. By this constellation it should
be possible to limit the responsible CMS. For breeding industry an application
of the markers given on the website allows to spare money and time in the
crossing work. The identification of male sterile genotypes enables a limitation
in the selection of pollen parents. It will further be of advantage, to
exchange the cytoplasms of these sterile mt-g
cultivars by the method of Spangenberg et al. (1991) or Rasmussen
et al. (2000).
Somatic fusion hybrids
In general the results have to be seen in front of a distinct nuclear
genome composition. They are not transferable completely to other fusion
populations with similar cytoplasmic combinations.
For confirmation of the effects found, additional fusion populations
have to be analyzed; on the other hand it is necessary to check their nuclear
genomes for deviations. Within the populations a small portion of
somaclonal variation could be proven with a list of probes which covered
the whole set of chromosomes. Due to the fact that the main part of mutations
have an unfit effect, there is a negative correlation of somaclonal variation
with yield parameters. But in none of the euploid hybrids the deviations
caused depressions in field performance (Lössl et al. 1994).
From this we can follow, that within the concerned hybrid population
the nuclear genomes are not affected from serious aberrations.
Interaction with the novel plastid-mitochondrial factors during accumulation
and degradation of polysaccharides constitutes the variable field performance
of individual hybrids.
The analysis of cytoplasmic organelle components within somatic hybrid
populations revealed high differences between the hybrids cp and mt genomes.
The skewed segregation ratio after regeneration phase of 1:6 in cp type
W and T was a first indication for an advantage of distinct cytoplasms
in fusion population Ia. This deviation from random plastid segregation
could be due to different frequencies of replication and organelle division
(Glimelius et al. 1981, Donaldson et al. 1994) and is indicative for a
higher performance of cp type T during in-vitro regeneration phase.
In the analyzed populations I, II, III, IV the segregated plastid genomes
did not show any recombinations and were highly conserved, whereas mitochondrial
genomes of the hybrids revealed novel DNA organizations R1 and R2. On first
view in population Ia somatic hybrids with cp type T had a significant
advantage with respect to starch content, but additionally an interaction
with the mitochondrial complement was observed by the comparison of recombinant
mt genomes. Horvath et al. (1992) suggests that recombinations are processes
of adaptation, which improve the nuclear-cytoplasmic interaction. The significant
difference in starch yield of recombination type R1 and R2 in dependence
of cp type W and T displays the association of distinct chondriome-plastome
arrangements with better performance levels.
For an evaluation of this effect mt type was predominantly mentioned
for Population Ia. One reason was, that population Ia had the lowest degree
of deviations in nuclear genomes. Hereby the risk was reduced, that differences
were due to protoclonal variation. Furthermore, in contrast to the other
populations only population Ia had a skewed cp type segregation. The skewed
ratio was associated with a high difference in starch content between the
two groups W and T, whereas the other populations show weak or low significant
correlations to starch content. All the populations Ia, Ib, Ic, IIc, IIIb,
IVa, IVb are obtained from different fusion parents. Population Ia
and Ib contain different nuclear genomes. Therefore it is supposed that
the effect, observed in population Ia is dependent on cytoplasmic interaction
with the nuclear composition. The other populations listed, show, that
there is in general a superiority of these clones, which segregated to
a higher number. If this relation seemed to be otherwise, than it turned
out, not to be rather significant.
In order to keep an overview, only the significant influences of organellar
configurations are stated. Homogeneous mt types had no influence on starch
production in population Ia. In order to confine mt regions responsible
for interactions with plastid type cpW/ cpT, the experiment focused on
mt recombination types R1 and R2. No other recombinations in potato mitochondrial
genomes have been described so far and mitochondrial types of other populations
did not show any significant correlations. Further evaluation of differentially
segregated mt types might reveal other interactions, in especially male
Distinct mt DNA conformations seem to influence mitochondrial performance
by their compatibilities with the plastid factors. In this respect it is
interesting, that plastid type T lacks five open reading frames which exist
in the 5´ region of the atpE locus of cp type W (Kawagoe and
Kikuta 1991). In maize Newton et al. (1990) showed that mitochondrial factors
may play a role also for chloroplast function when they found a mutant
mt gene (coxII) which caused an abnormal function of photosystem I. Only
an optimized mt-cp configuration allowed an efficient interaction with
the new generated cell composition.
It is a question, whether selection pressure works on the level of
sugar content in the heterotrophic culture medium. Sugar content in our
regeneration media is reduced to one third of the normal concentration
in MS medium as a prevention of vitrification effects. This might be a
selection pressure, but the bottleneck for cytoplasm performance could
also be located in other metabolisms. It is supposed that different compatibility
of distinct regions of cp and mt genome is the reason for cytoplasmic interaction:
The rare organellar configurations apparently undergo basic problems, reducing
vitality during replication of organelles. This could be any missing recognition
of regulatory factors encoded by the nucleus, which affects transcript
editing, splicing, processing and translation processes. The deviation
concerning the duplicated atp6 copy in Population IVa is indicative for
an abnormal mt genome replication, which contributes to the superiority
of homogenous mitochondrial genomes (Lössl et al. 1994). Such
differences could be due to ´biological costs´, which are associated
to nuclear-mitochondrial incompatibilities as reported by McVetty and Pinnisch
(1994) for different plasma types in Brassica.
In starch content hybrids with an additional atp6 copy yielded
significantly lower than hybrids without this deviation. In this population
deviant expression of organellar genomes could be a reason for incompatibilities
between cellular compartments. A possible source of variation in this hybrid
population is the presence of additional mitochondrial proteins (Lössl
et al. 1999). Such deviations in the cytoplasmic expression often are associated
with male sterility as a result of mt genome recombinations (Leaver et
al. 1988, Köhler et al 1991, Horn and Friedt 1999). The reduction
in fertility and vitality is in accordance with the observation quoted
above, that hybrids affected from extreme recombinations exhibited lower
vigor than hybrids with homogeneous chondriomes.
In this respect it will be of relevance to analyze the mt types on
their expression level and to find the reasons for the improved nuclear
Thus, cytoplasmic analysis is expected to deliver informations for
somatic genetics and is involved in polygenic breeding aims, which are
dependent on an optimized nuclear cytoplasmic compatibility. Novel mt-cp-configurations
generated by cell fusion may be an enrichment for breeding not only in
vegetatively grown plants like potato but also in sexually propagated crops.
|Figure 1a,b,c. Cytoplasmic
types detected by PCR primers. Lanes: 1) Ponto, 2) Adretta, 3) Karlena,
4) Desiree, 5) Sieglinde, 6) Cilena, 7) Assia, 8) Heidrun, 9) Helios, 10).
DNA-Standard. DNA product sizes resulting from different PCR primer combinations
are given in Table1.
ALC_1 and ALC_3
differ between cp types T and
Lambda Standard (St.) cut with HindIII
ALM_4 and ALM_5
differ between mt types a,
b and g.
Lambda Standard (St.) cut with Sty
ALM_6 and ALM_7
differ between mt types
a or b and
Lambda Standard (St.) cut with HindIII.
||Figure 2. Plastid-mitochondrial segregation
in population Ia. The mitochondrial genomes of the fusion population consisted
of clones with R1 and R2 and homogeneous mt genomes (a/b).
The latter subordinated to the main effect of plastids in population Ia,
given in Table 4. Configurations R1 and R2 deviated
from this effect and therefore were analyzed on further interactions. The
ratio is the absolute number of clones simultaneously.
||Figure 3. Interdependence
between cp- and mt type R1 or R2 of a somatic hybrid population in
starch yield (kg/plot). Fusion hybrids of population Ia were determined
as groups "R1" and "R2" and subgrouped according to their plastid types
T or W. Given are the 95% confidence intervals for starch yield [kg] of
the four groups for four years.
|Table 1. Primer pairs
used for identification of cytoplasmic type. The table gives the
primer name, the sequence, the primed region and the product size, specific
for W/a, W/g
and T/b. ALC primers are used for determination
of plastid types, ALM primers for mitochondrial types.
|Table 2. Frequencies of Plastid-Mitochondrial-Types
within the German seed board and haploids. T identifies tuberosum type,
S and W are wild types
Percentage in german assortment
|Table 3. Correlations of effects of different
cytoplasms, observed within the spectre of cultivars, in di-haploids and
a reciprocal F1 population. Probing was carried out according to Lössl
et al. (1999). Replications mean the number of different cultivars and
respectively the number of dihaploid clones, which were compared in groups
of identical cytoplasm types. Cultivar experiments yielded similar results
as published by the official „BundesSortenamt“.
> Wa Tb
: 46 + 79 ]
P = 0,01
|| Wa Tb
||primers: ALM_4/ 5
> Wa Tb
: 46 + 79 ]
|P = 0,01
|| Wa Tb
||primers: ALM_4/ 5
> Wa Tb
: 46 + 79 ]
|P = 0,05
| Wa Tb
| Wa Wg
||P = 0,01
| Wa x Wd,
Wa x Wa
||P = 0,01
|Table 4 Correlations of different cytoplasmic
configurations with starch content, observed within fusion hybrid populations.
Probing was carried out according to Lössl et al. (1999).
Wa (+) Tb
Wa (+) Tb
Wa (+) Tb
Wa (+) Se
Wa (+) Wg
Wa (+) Wd
Wa (+) Se
.cpT > cpW
[270 : 45]
[68 : 42]
cpW = cpT
[5 : 30]
cpS > cpW
[10 : 54]
a > g
[65 : 20]
b > d
[35 : 10 : 5]
[317 : 29]
P = 0,01
P = 0,05
P = 0,05
P = 0,05
P = 0,01
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The authors are grateful to Mr. A. Barth, Mrs. E. Gerick and
C. Kornbauer for technical assistance.
This work was supported by grants of the BMBF under the number 0310768.
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