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Hybrid potato breeding for improved varieties



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© Burleigh Dodds Science Publishing Limited, 2016. All rights reserved.
Hybrid potato breeding for improved
Pim Lindhout, Michiel de Vries, Menno ter Maat, Su Ying, Marcela Viquez-Zamora and
Sjaak van Heusden, Solynta, the Netherlands
1 Introduction
2 The scientifi c basis for hybrid potato breeding
3 The state of the art of hybrid potato breeding
4 Production of and commercialization of hybrid seed cultivars
5 Inbred lines for genetic research
6 Cropping systems based on true seeds
7 Case studies
8 Conclusion
9 Where to look for further information
10 Acknowledgements
11 References
1 Introduction
The cultivated potato, Solanum tuberosum, can be reproduced generatively through
seeds and vegetatively through tubers. This may have evolutionary advantages: seeds
may provide better survival under extreme conditions, such as frost or drought, and can
remain viable in the soil for years. When conditions are mild, tubers survive in a dormant
state for a couple of months. When conditions become favourable again, their fast and
strong sprouting provides a clear competitive advantage over other plants in the same
ecological niche.
In traditional potato breeding, each breeding cycle starts with a cross between
two genotypes, usually tetraploid varieties, followed by many years of selection and
multiplication (see Chapters 2 and 3). The advantage of this approach is uniformity: the
tubers are clones and thus genetically identical. The disadvantage is the low genetic gain in
each lengthy breeding cycle, as the genetic composition of the two parental genotypes is
just reshuffl ed, including alleles which negatively affect plant growth and development. As
a result, potato yield has not signifi cantly been improved over the past century (Douches
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2 Hybrid potato breeding for improved varieties
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et al., 1996; Vos et al., 2015). In addition, the reproduction of seed tubers is less than a
factor 10 per season. It takes many years to build up suffi cient quantities of seed tubers for
commercial production, and the risk of contamination by pathogens increases with each
multiplication step.
True potato seed (TPS) has been promoted as an alternative for seed tubers because
TPS is easy to store and devoid of most soil-borne pathogens. In South Asia, East Africa
and the Andes, TPS is used mainly by subsistent farmers (Almekinders et al., 1996). TPS is
produced by crossing parent plants that have been selected to produce a hybrid variety.
The parents are propagated vegetatively, similar to seed tuber propagation. As the parents
of a TPS variety are heterozygous, all seeds of a TPS cultivar are genetically different. This
results in a highly variable crop that is not acceptable in most markets, such as the high
value markets of Europe and North America.
Since the success of hybrid breeding in corn in the 1930s, breeders have adapted a
hybrid breeding system for many crops (Fig. 1; Crow, 1998; Troyer, 2006; Hua et al., 2003).
Typically, hybrid cultivars produce higher yields and show high crop uniformity (Rijk et al.,
2013). In addition, the breeding system is fast and effi cient and new traits can rapidly be
introduced by marker-assisted introgression.
These advantages are also expected for potato: hybrid potato varieties will be higher
yielding, will need less crop protection chemicals due to disease resistance and will have
better quality for processors and consumers (FAO et al., 2015). A hybrid breeding system
for potato offers two additional advantages: fast multiplication of hybrid seeds and easier
logistics, as clean true seeds can easily be produced, transported and stored (Duvick et al.,
Figure 1 Fivefold increase in corn yields since the introduction of hybrids (Troyer, 2006).
check if ‘is
less than
a factor
10 per
season’ is
OK as is.
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Hybrid potato breeding for improved varieties 3
However, self-incompatibility and inbreeding depression have for long hindered
progress in hybrid potato breeding (De Jong and Rowe, 1971; Charlesworth and Willis,
2009; Jansky et al., 2016). These two limitations have recently been overcome by
introducing a self-compatibility restorer gene and by large-scale and consistent breeding
(Lindhout et al., 2011a).
This chapter describes the scientifi c principles and applied aspects of hybrid potato
breeding. The successful introduction of the principle of hybrid potato breeding was
described in 2011 (Lindhout et al., 2011a). Since then, we have focused on further
developing the potato hybrid breeding system, especially genetic studies to establish
a genetics-driven hybrid breeding system. In a recent paper, Jansky et al. (2016) have
confi rmed the possibilities of such approach.
This is the fi rst publication on the state of the art of a hybrid potato breeding
programme. The authors, all working at Solynta, want to emphasize that scientifi c papers
on this topic are not available yet, and hence we have to rely entirely on the results of
the Solynta breeding and research programme. Still, in presenting these results, we hope
to contribute to a better understanding of the principles and applied aspects of hybrid
potato breeding.
2 The scientifi c basis for hybrid potato breeding
2.1 The principle of hybrid breeding
The basic idea of hybrid breeding is to combine the genes of two parent genotypes,
both of which may harbour suboptimal alleles, resulting in weaker performance. If parents
have different suboptimal alleles, hybrid offspring can show increased vigour and yield,
designated heterosis , as the suboptimal alleles in one parent may be compensated by
the favourable genes from the other parent (Birchler et al., 2010; Gopal, 2014; Fig. 6). If
the parents are completely homozygous, the resulting hybrid offspring will be partially
heterozygous and genetically uniform. By testing many hybrid offspring under relevant
cultivation conditions, the best combining parents are identifi ed. These are maintained
and propagated in separate groups as heterotic pools for further breeding (Brown and
Cagliari, 2011).
Thus, hybrid breeding has two distinct processes: development of homozygous parent
lines and production and testing of experimental hybrids.
2.2 Diploids are more effi cient than tetraploids for hybrid
Homozygous diploids are faster to generate than homozygous tetraploids. For instance,
seven generations of selfi ng are required to obtain 50% homozygous loci starting from a
tetragenic tetraploid heterozygote (carrying four different alleles). The same homozygosity
level is reached starting from a heterozygous diploid by only one generation of selfi ng
(Haldane, 1930; Fig. 1). For this reason, hybrid potato breeding is more effi cient at the
diploid level.
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4 Hybrid potato breeding for improved varieties
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2.3 Generation of homozygous diploid potato genotypes via
haploidization and via inbreeding
Haploid genotypes can be generated from an egg cell (gynogenesis) or from pollen,
often via anther culture (androgenesis). In potato, some haploids have been generated
by anther cultures. The resulting haploids were converted into diploids by chromosome
doubling. However, the resulting homozygous diploids were very weak and sterile (van
Breukelen et al., 1977; Uijtewaal et al., 1987b), hampering their usage in breeding.
Haploidization has been more successful in crossable species like S. chacoense
(Cappadocia, 1990; Phumichae et al., 2005; Phumichae and Hosaka, 2006) and S. phureja
(Chani et al., 2000).
A reason for the failure to produce vigorous doubled haploids may be the transition
to complete homozygosity in one step. Inbreeding depression may be so severe that
homozygous plants are too weak to survive. Repeated selfi ng, on the other hand, might
lead to a more gradual improvement of homozygosity. However, in potato, inbreeding is
seriously limited by self-incompatibility, which prevents self-fertilization. Still some rare
examples of homozygous diploid plants have been generated by inbreeding but again the
homozygous diploid plants always showed a strong inbreeding depression, which limited
their usage in breeding (De Jong and Rowe, 1971; Charlesworth and Willis, 2009).
2.4 Large genetic variation in potato causes inbreeding
The tetraploid and outcrossing nature of commercial potato is likely responsible for the
large genetic variation. In a study on the allelic composition of 800 genes in 83 potato
cultivars, an average frequency of 3,2 alleles per locus within a genotype was identifi ed
(Uitdewilligen et al., 2013). Among the 83 cultivars, often more than ten alleles per locus
homozygocity level
generations of selfing
diploid AB
tetraploid AAAB
(digenic simplex)
tetraploid AABB
(digenic duplex)
tetraploid AABC
tetraploid ABCD
Figure 2 The theoretical increase in homozygosity in diploids and tetraploids through inbreeding,
adjusted from Haldane et al. (1930).
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Hybrid potato breeding for improved varieties 5
were observed. In addition, the frequency of single nucleotide polymorphisms (SNPs)
in potato is 1 in 15 30 base pairs (PGCS, 2011; Visser et al., 2014). This means that
the genetic distance between the two sets of chromosomes within one diploid potato
genotype is four times larger than the genetic distance between the genomes of man and
chimpanzee (CSAC, 2005).
The high frequency of allelic variation has the inevitable consequence that weak alleles
that have a negative effect on plant fi tness are maintained. Such alleles remain hidden in
the large buffer of four genomes, but reveal themselves upon inbreeding when the chance
for homozygosity increases. This is even more manifested at the diploid level where the
homozygosity level more rapidly increases upon inbreeding (Fig. 2).
This large genetic variation is also helpful for breeding as it forms a genetic reservoir of
useful genes. It is a challenge to identify alleles that contribute most to plant performance.
As potato has 39 000 genes, the identifi cation and usage of the best alleles per locus,
including interactions between them (epistasis), will gradually take place over decades of
research and breeding (PGSC, 2011). Corn may serve as a good example, whereby, after
a century of dedicated breeding by numerous breeders worldwide, a genetic gain of over
1% per year is still achieved (Troyer, 2006).
2.5 Crossable diploid species and tetraploids increase genetic
reservoir for diploid breeding
The potato germplasm available for breeding comprises many species, including diploid
species (Jansky and Peloquin, 2006). These have been used as source to introduce
resistance genes into cultivated germplasm. Breeders often use diploid potato to rapidly
combine favourable traits that can be introduced into the tetraploid germplasm by direct
crossings, bridge crossings or via chromosome doubling (De Mainea, 1982; Chauvin et al.,
2003). The diploid breeding programme at Wageningen University (Hutten, 1994) has
generated donor lines that harbour the most important traits for potato breeding (Table 1).
Additional diploid germplasm is available from potato gene banks and public research
institutes such as University of Wisconsin-Madison, United States; Potato Germplasm
Enhancement Laboratory, Japan; Gene Bank at Gatersleben, Germany and International
Potato Centre, Peru.
Another source of diploid germplasm is tetraploids that can be prickle pollinated to
generate diploid offspring, designated dihaploids (Uijtewaal et al., 1987a). A collection of
dihaploids obtained from one tetraploid harbours the full set of genes from the tetraploid
and can be exploited in a diploid potato breeding programme.
In conclusion, the large genetic variation in potato and in its wild relatives, combined
with the technology to switch between ploidy levels, provides a tremendous wealth of
germplasm available for diploid hybrid breeding.
2.6 Diploid potato may perform equal to tetraploids
Most important food crops such as rice, corn and soybean are diploid. Sugar beet cultivars
were initially tetraploid, then triploid and since 2000, all new cultivars are diploid. In potato,
it has long been assumed that tetraploids outperform diploids (Rowe, 1967; Hutten et al.,
1994). Occasional observations have contradicted this assumption: Progeny of diploid
potato USW4 with S. chacoense M6 produced large tubers and high yield (Lipman and
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6 Hybrid potato breeding for improved varieties
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Zamir, 2007; Jansky et al., 2014). Uijtewaal et al. (1987b) observed that heterozygous
diploid potato outperformed all homozygous di- and tetraploid derivatives. These
results from potato and from other crops support the expectation that diploid potato will
eventually replace tetraploid potato for commercial usage.
3 The state of the art of hybrid potato breeding
In 2008, Solynta started its research by making crosses between diploid potato germplasm,
obtained from a pre-breeding programme from Wageningen University (Rutten, 1994),
and a homozygous accession of the wild species S. chacoense , carrying the dominant self-
compatibility controller gene Sli (Hosaka and Hanneman, 1998a,b; Phumichai et al., 2005;
Lindhout et al., 2011a). The F
1 plants were extremely vigorous and about half of them
produced many berries upon self-pollination. This was considered a major breakthrough
as these were, to our knowledge, the fi rst vigorous, self-compatible diploid potato plants
ever obtained. These F
1 plants were highly heterozygous. The fi rst generation after selfi ng
(designated F
2 ) should harbour at least 50% homozygous loci. As many of these loci might
Table 1 The Sli -gene donor, designated DS and 16 diploid potato
germplasm, designated D1 D16, used for hybrid breeding at
Solynta. The trait abbreviations are according to Hutten (1994)
Abbreviation Short description
DS Sli- gene donor
D1 Early (maturity), long, Y , Qcook
D2 Early, Y, Qcook
D3 R3 , H1 , Gpa2 , RXadg , Y (yellow fl esh)
D4 Grp1, early, long (shape), Ro1 ( H1 ?)
D5 Early, long, Y , Qfry
D6 Long, Y , Qfry, H1 , Qcook
D7 Early, long, Y , H1 , Qcook, Zep (orange fl esh)
D8 Early, y (white fl esh)
D9 Qstarch, Y
D10 Wild species hybrid: phyt avl
D11 Wild species hybrid: phyt rch
D12 Round (shape), Qcook, Qfry
D13 Early, round, Zep , Y , Spectacled, Qcook, blue
D14 Wild species hybrid: phyt tar
D15 Wild species BC1: early, phyt vnt1 , round, Y , H1
D16 Early, round, y (white fl esh)
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Hybrid potato breeding for improved varieties 7
harbour weak alleles , it was expected that F
2 plants would be too weak to survive. We
also made BC
1 populations by backcrossing to the S. chacoense parent.
As expected, the F
2 and BC
1 showed weak growth and many died in the fi eld. However, a
number of plants survived and 10% of the surviving F
2 plants even proved self-compatible
(Lindhout et al., 2011a). We generated the second inbred generation, designated F
3 and
made crosses between self-compatible individual F
3 plants. These inbred plants were
tested with SNP markers to confi rm their genetic identity as real inbreds. These results
indicated that breeding hybrid potato was now feasible (Lindhout et al., 2011b).
The weak plant performance, the poor tuber quality and the low yield of the inbreds
was not only due to inbreeding depression, but was also caused by the wild S. chacoense,
a species that hardly produces tubers. So, by this approach, we not only started a hybrid
potato breeding system, but also the process of domestication of a new diploid potato
based on an interspecifi c cross of diploid S. tuberosum and the wild species S. chacoense.
We hypothesized that developing vigorous inbred lines is challenging as for each of the
39 000 loci the most favourable alleles should be identifi ed and combined. Unfavourable
genes with large effects on plant performance are identifi ed easily and hence selecting
increased plant performance is initially easy and fast. Undesired characters from the
S. chacoense parent such as abundant stolons, small leaves and twisted, small and low-
yielding tubers were removed in a few breeding generations.
Sli is a dominant gene (Phumichai et al., 2005; Phumichai and Hosaka, 2006). However,
the successful expression of this gene requires a vigorous plant that is fertile and supports
self-pollination. Often these criteria are not met. Therefore, the frequency of self-
compatible plants is usually lower than expected based on a monogenic trait. Inbreeding
depression is exhibited as weaker plant growth upon higher generations of inbreeding
(Fig. 3). As a consequence, the self-compatibility level tends to decrease upon further
selfi ngs. The fi rst inbred lines obtained by Solynta, containing over 95% homozygous
Figure 3 Inbreeding in the diploid Solynta germplasm in winter 2014 15. The Fx indicates x-1 generation
of selfi ng after the last cross was made. Data are averaged over the complete trial consisting of over
5000 plants. The scale of plant vigour ranges from 1 = very weak via 3 = average to 5 = very strong.
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8 Hybrid potato breeding for improved varieties
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loci, hardly produced progenies and the seedlings were extremely weak. Therefore, new
crosses were made between F
3 and F
5 inbred lines and selfi ng was started again from
these F
1 s to continue selecting parent lines that combine benefi cial traits. As a result,
performance of the inbreds improved over the following generations (Fig. 4).
Lady Roset ta
Lady Ana
Hybrid A
Hybrid B
Hybrid C
Hybrid D
Hybrid E
Hybrid F
Hybrid G
Hybrid H
Inbred B
Inbred C
Inbred D
Inbred E
Inbred F
Inbred G
Inbred H
Tuber Yield (gr/plant)
LSD α=0,05
Figure 5 Hybrid performance of the fi rst diploid experimental potato hybrids. Seed tubers were
harvested from greenhouse-grown plants, raised in the winter from tubers (checks) and seedlings
(hybrids and inbreds). The seed tubers were planted on 8 May 2015 in a trial fi eld on sandy soil in
Wageningen and harvested on 17 September 2015.
Figure 4 Examples of Solynta diploid germplasm. Plants were raised from seedlings and grown in the
greenhouse in the summer season of 2014 (left panel) and 2015 (middle and right panel). They were
among the best genotypes in a breeding programme comprising over 15 000 plants.
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Hybrid potato breeding for improved varieties 9
In mature hybrid breeding systems, parents of the hybrids are selected based on general
combining abilities (GCA), whereby molecular markers can be used to better predict the
breeding value of the parents (Tobias et al., 2009). As no historic data were available on
combining abilities of diploid potato parents, the selection of the fi rst parents was based
on the performance of the parents themselves.
Following the predictions of good performing inbred lines, crosses between these lines
were made resulting in 45 potato hybrids which were tested in the fi eld together with 20
inbred lines in two replicates of two plants per plot. The yields varied from 83 to 580 g/
plant (Fig. 5). Thirteen hybrids scored higher than any of the inbreds, also outperforming
the check variety Lady Anna, while some also showed a strong heterosis for yield (Fig. 6).
The trials are being repeated in 2016 by using seed tubers raised from the fi eld.
In addition, a new series of 216 experimental hybrids was generated in the winter-season
2014 15, mainly based on F
6 parent lines. The hybrid seeds were sown immediately after
harvest and ten seedlings per hybrid were transplanted into the fi eld in June, 2 months
later than the usual seed tuber plantings. Still the yield of some hybrids was higher than
500 g/plant and the tubers were similar in size and shape to commercial seed tubers.
The results of the fi rst experimental hybrids illustrate the potential of diploid hybrid
potato varieties. As the fi rst hybrids were randomly made without any a prior information
about the combining abilities of the parents, it is expected that the next series of hybrids
based on the results of these fi eld trials will perform better and may show a stronger
overlap with commercial controls.
4 Production of and commercialization of hybrid seed
The production of hybrid potato seeds is mainly done by hand pollinations. The seed
yield per plant varies from hundreds to many thousands of seeds. Each successful hand
pollination generates a berry with 50 150 seeds and each plant produces 5 50 berries. This
Figure 6 Example of heterosis in diploid hybrid potato. The plants were from the same trial as in
Figure 5. At the left the female F
3 parent and at the right the male F
5 parent, while the hybrid is in the
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is similar to other vegetable Solanum crops, like tomato and pepper, where commercial
seed is produced by hand pollination.
The emasculation of the fl owers is a time-consuming part of the hand pollinations.
Male sterility systems have been reported that make emasculations redundant and hence
reduce seed production costs (Li, 2008). Alternatively, functional male sterility may be
used, whereby pollen is prevented to land on the stigma of the same fl ower as the exerted
stigma may be manually pollinated by pollen of the male parent before the own pollen
may reach the stigma (L ö ssl et al., 2000; Abrol et al., 2012).
The transport of potato seeds over the world is very restricted. This is because the
dominant breeding systems are based on the production of seed tubers and hardly any
rules are in place for potato seeds. As a consequence, most countries consider potato
seeds as belonging to the highest risk classes. This is remarkable as seed tubers may
harbour any of over 200 species of pathogens that attack potato (Delleman et al., 2004).
In contrast, only six pathogens are seed-borne (Solomon-Blackburn and Barker, 2001).
These are fi ve viruses and a viroid, which are absent in major potato-growing regions like
the Netherlands. So, potato seeds are very safe and regulations will need to be adjusted
The registration process for breeders rights poses a similar situation: in countries
which are members of the International Union for the Protection of new Varieties of Plants
(UPOV), the path to commercialization of a hybrid starts with the registration process to
obtain breeders rights, which is based on seed tubers. So, protection by breeders rights
of hybrid potato seed cultivars is not yet possible. The European Union (EU) is adjusting
the legislation process, but it may still take several years before this is established. Other
non-UPOV countries will likely follow later.
5 Inbred lines for genetic research
Inbred lines allow the generation of mapping populations such as F
2 , BC
1 and BC
2 . As
the parents have limited allelic variation, the signal-to-noise ratio is much higher than
in studies with heterozygous tetraploid populations. Moreover, putative quantitative trait
loci (QTL) can effectively be confi rmed in dedicated populations that are selected to
segregate for the loci under investigation, and are fi xed for other regions on the genome
(Wang et al., 2008; Schmalenbach and Pillen, 2009; Fu et al., 2010). In addition, new
genetic populations can be generated, which are very powerful for quantitative genetic
studies, like nearly isogenic lines (NILs), recombinant inbred lines (RILs) and libraries of
introgression lines (Young et al., 1988; Paran et al., 1995; Jeuken and Lindhout, 2004;
Finkers et al., 2007; Zhang et al., 2005; Chen et al., 2010; Viquez et al., 2014).
Genetic studies in potato have been done at the tetraploid level and at the diploid level.
Tetraploids may support simple genetics like the mapping of resistance genes (Solomon-
Blackburn and Barker, 2001), but quantitative studies are less reliable as the genetic
noise of the numerous highly heterozygous loci is high (unexplained error). Genome-
wide association studies (GWAS) at the tetraploid level will always generate hundreds of
potential leads, but only a few hits may be meaningful (Li et al., 2010; D Hoop et al., 2014).
More accurate and reliable quantitative studies were done at the diploid level, initially by
crossing heterozygous parents (Prasher et al., 2014) and, more recently, in a diploid F
population (Endelman and Jansky, 2016).
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Hybrid potato breeding for improved varieties 11
The possibility of using self-compatible, homozygous inbred lines for genetic studies
offers three powerful advantages:
1 Only one allele per homozygous locus is present.
2 Only two alleles per heterozygous locus are present.
3 Backcrosses and selfi ngs are feasible for confi rmation studies.
For more than fi ve decades, mutant studies have uncovered new alleles of important
genes and have helped to confi rm or determine the function of genes. Although advanced
technologies like the CRISPR/Cas system (Belhaj et al., 2013) are likely to replace the
methods by which mutants are made, mutants will remain powerful tools to discover
unknown phenotypic traits or to study induced alleles that also have the advantage to be
free of deregulation rules.
5.1 The fi rst completely homozygous self-compatible diploid
inbred line in potato
Most Solanum species that are crossable with cultivated potato are self-incompatible. An
exception is S. chacoense (Hosaka and Hanneman, 1998a; Hawkes, 1990; Jansky et al.,
2014) and introducing the Sli- gene from S. chacoense into cultivated diploid potato
resulted in fertile self-compatible offspring (Lindhout et al., 2011a). After several rounds
of inbreeding, highly homozygous self-compatible inbreds were generated. The level of
homozygosity was assessed by using SNP markers to investigate the effects of inbreeding
(Fig. 7). A strong correlation was observed between the overall level of homozygosity and
reduced self-compatibility. By new series of crosses, selections and selfi ngs, the agronomic
Homozygosity percentage
Plants of research lines
Homozygosity level of four research lines
Figure 7 The increase of the level of heterozygosity in the offspring (BC
1 F
7 ) of four BC
1 F
6 plants
obtained by selfi ng of the BC
1 (D1 × F
1 [DS × D1]) TQ: Please check the usage of brackets is as per
style. (Table 1). The width of the bar indicates the number of plants in the progeny within the class with
plants of the same homozygosity level. The calculations were based on a total of 150 markers.
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12 Hybrid potato breeding for improved varieties
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performance of the inbred lines continuously improved. After six generations of selfi ng,
a homozygous self-compatible line was generated with only 1 out of 150 SNP markers
still heterozygous (Fig. 8). Genotyping by sequencing techniques make it now possible to
check the level of homozygosity in more detail.
5.2 Genetic studies in segregating diploid F
2 populations
Recently, Endelman and Jansky (2016) published the fi rst results of a mapping study in an F
population of diploid potato. This was based on a cross between the doubled monoploid
potato DM1-3 and M6, which is an S
7 inbred line derived from the self-compatible wild
relative S. chacoense . A single F
1 plant was then self-pollinated and an F
2 population of
109 genotypes was grown, genotyped (>2200 SNPs) and phenotyped. Meijer et al. (2016)
analysed an F
2 population (108 markers) based on a cross between two clones, namely DS
(a homozygous diploid S. chacoense clone containing the self-incompatibility overcoming
Sli- gene) and D2 (a partly heterozygous diploid S. tuberosum clone; see also Table 1).
The results of both studies are comparable: tuber shape is associated with a region on
chromosome 10, fl esh colour with a region on chromosome 3 and tuber and pigment colour
on chromosomes 2, 10 and 11. These QTLs were identifi ed at the same loci as described
in literature (van Eck et al., 1993, 1994). In both studies, additional QTLs were identifi ed.
Furthermore, there is an overwhelming reservoir of potential useful QTLs in the potato
germplasm (Bradshaw et al., 2007) and thus also in the dihaploids that can be made. Such
QTLs can now be more reliably studied at the diploid level and this will ultimately lead
to the identifi cation of the underlying genes. A limited subset of the diploid germplasm
may already harbour many important traits for potato breeding (Table 1). Relevant genes
01 02 03 04 05 06 07 08 09 10
11 12
Figure 8 The fi rst essentially homozygous self-compatible diploid potato. The position of the SNP
markers is based on the published sequence (PSGC, 2011). The 12 vertical bars indicate the 12
chromosomes. Red bars indicate homozygous D1, blue bars indicate homozygous DS (Table 1). The
green line on chromosome 11 indicates a heterozygous scored marker.
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Hybrid potato breeding for improved varieties 13
for these traits can be introgressed into vigorous and fertile diploid genotypes to develop
progenies with maximum genetic fi xation to minimize genetic noise. Such genotypes are
unique materials for further detailed genetic studies.
5.3 Marker-assisted backcrossing
Another application for inbred lines in potato is introgression breeding. This requires
knowledge of diagnostic markers for the gene of interest, preferably inside the gene,
markers for the recurrent parent genome and a self-compatible homozygous backcrossing
parent. There are dozens of well-studied resistance genes in potato that can be used
for introgression breeding. This paves the way for a marker-assisted backcrossing (MAB)
programme in potato (Frisch and Melchinger, 2005).
To this end, homozygous inbred lines are crossed with a diploid donor carrying a specifi c
gene of interest. In two backcrosses, NILs can be generated by selection with diagnostic
markers for the gene of interest and against markers in its fl anking regions, combined
with selection for markers well distributed over the potato genome (whole background
selection). Such NILs can harbour over 98% of the recurrent genome in combination with
the specifi c gene. Both parents of a hybrid may have an introgressed gene, resulting in a
double stack hybrid. To introgress specifi c genes in a homozygous parental line will take
2 3 years. These MAB programmes are routinely used in other crops and are also feasible
in potato (Mallick et al., 2015; Jeong et al., 2015).
6 Cropping systems based on true seeds
The production of commercial tubers in most parts of the world starts with seed tubers.
These have a large reservoir of nutrients for the growing shoots, allow a rapid initial plant
growth and fast leaf coverage of the soil, which is one of the most critical factors for potato
yield. In contrast, potato seeds are extremely tiny, about 2500 seeds per gram. As a result,
during the fi rst period after germination the young seedlings are very vulnerable for abiotic
stresses like drought, frost and heat. Field emergence has been reported between 50%
and 80% with acceptable tuber yields under different experimental conditions (El-Bedewy
et al., 1994; Renia, 1995). However, without a protective environment, the risk is very high
that an emerging seedling will not survive, even when it is pelleted or primed.
This chapter describes alternative strategies to circumvent the exposure of week
seedlings to harsh conditions.
6.1 Production of seedling tubers in greenhouse
Seedling tubers can be produced under greenhouse conditions by sowing in a medium
with suffi cient water supply and at optimum germination temperature of 15 20°C (Struik
and Wiersema, 2012). As soon as seedlings reach 5 10 cm in length, they are transplanted
in pots. The desired tuber size, the available space in the greenhouse and the length of
the growing period will determine the pot size, nutrient supply, day/night temperature and
light regime (for a detailed protocol see Struik and Wiersema, 2012). Densities of 80 170
2 are common in greenhouses (Lommen, 1995; Tierno et al., 2014). There is a
trade-off between number of tubers, size of tubers, planting density and time to harvest.
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Depending on the exact conditions, end-product requirements and production costs,
the optimal combination is chosen. Alternatively, hydroponic techniques are available,
whereby the roots are grown in a dark, humid and soilless environment in two layers, one
for nutrient uptake and the other layer for producing mini-tubers (Lommen, 2007). The
seedling tubers are picked at regular intervals and the total yield per plant may reach
dozens of tubers. The soilless culture assures clean seed tubers. The conditions and
picking regime are set to have optimal numbers and tuber sizes.
6.2 Production of seedlings for commercial crop
Greenhouse-grown seedlings can also be used to start the cultivation of a commercial
crop. This system is equivalent to the one used for lettuce, leek, cabbage and onion
(Leskovar et al., 2014). Technically, commercial potato production from seedlings is
feasible and maximizes the benefi ts of true seeds. When potato transplants are grown
as a ware potato crop, a whole new cultivation system must be developed. Important
elements are: transplanting systems, plant spacing including ridging or bedding, the use
of soil coverage, weed control, irrigation and harvesting methods. Further mechanization
and dedicated cultivation systems will be optimized for cropping systems that start with
potato seedlings (Roy et al., 2015). In Kenya, tuber yields of 30 tonnes/ha were obtained,
whereby seedling transplants were used as starting material for a commercial cultivation
(Muthoni et al., 2014). This already represented 50% higher yield than average in Kenya
(Wang om and van Dijk, 2013).
6.3 Seedling tubers as starting materials for a commercial crop
Seedling tubers are equivalent to mini-tubers that are produced from tissue culture,
which is routinely done to start a new multiplication round with clean basic seeds (Amin
et al., 2014). They are certifi ed as G1 material. The great advantage of seedlings are the
reduced costs, compared to in vitro grown plants, and the fl exibility to start the production
whenever and wherever needed, as seeds can easily be stored and transported. As the
cost to produce mini-tubers from in vitro grown plants is very high, in the present potato
system at least three rounds of fi eld multiplications are needed to dilute these high costs
over many seed tubers. For a hybrid seed system, the cost of producing seedling tubers
is much lower and hence fewer propagation rounds are needed. Therefore, seedling
tubers should be multiplied only 1 year and then released to commercial farmers. Such a
system also fi ts better to the fast introduction of new cultivars, which is typical for a hybrid
breeding system.
6.4 Production of seed tubers from seedlings in fi eld
At present, the production of seedling tubers is mostly done under tropical conditions.
Seeds are sown in a simple greenhouse or in the fi eld under plastic cover with plant
densities of 80 100 plants/m
2 (Kumar, 2014; Struik and Wiersema, 2012; Fig. 9). When the
seedlings have reached 5 10 cm in length, they are transplanted to the fi eld, in ridges,
at a defi ned plant density. Additional hilling will increase the number of seed tubers per
plant (Wiersema, 1986). To decrease the risk of root damage, the complete substrate is
transplanted with the seedling. Plantlets need some time to adapt to outdoor conditions
before transplanting directly to the soil (Gopal, 2004). In South Asia, transplants are
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Hybrid potato breeding for improved varieties 15
planted on the sides of the ridges to benefi t from the shade and higher soil humidity. The
planting distance may be adjusted to compensate for a shorter growing season compared
to seed tubers, if appropriate. The cultivation conditions are similar to traditional potato
cultivation systems. In Egypt, seed tuber yields of 40 60 tonnes/ha were obtained in such
a system, based on tetraploid TPS populations (El-Yazied et al., 2004).
Seedlings are more sensitive to frost and drought than seed tubers. Thus, transplanting
is done in a frost-free season and with irrigation. Compared to the traditional systems of
producing seed tubers, whereby the tubers are planted far before the last night with frost,
the length of the growing season of transplants may be 2 months shorter. In addition, the
plant development may be further delayed due to a transplanting shock and weak initial
growth. Hence, tuber numbers are lower and tuber sizes are smaller compared to seed
tuber grown plants, causing severe yield reductions. Plant density may be increased to
compensate for these reduced yields per plants.
7 Case studies
7.1 Combatting Phytophthora infestans
Late blight, caused by the oomycete Phytophthora infestans , was responsible for the Irish
famine of 1845 47 (Fry, 2008). All potato cultivars were susceptible to the disease and
suffered severe yield losses that led to food shortage. Since then, breeders have selected
cultivars with fi eld resistance and from the early twentieth century onwards, have introduced
specifi c R -genes, often sourced from wild relatives. However, cultivar Pentland Dell, which
Figure 9 The fi rst seedlings of the fi rst diploid hybrid potato hybrids in Democratic Republic of Congo.
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16 Hybrid potato breeding for improved varieties
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carries three R- genes, already turned out susceptible to a new virulent race of the fungus
even before its widespread use and just 4 years after its introduction (Malcomsum, 1969).
At present, the optimal way to control P. infestans is a combined approach of clean seed
tubers, clean soils, early and preventive chemical protection and destruction of crop debris
after harvest. The global annual cost of cultivation measures and yield loss is estimated at
US$5 billion (Duncan, 1999).
P. infestans reproduces mainly clonally. With hundreds of billions of spores produced per
hectare in an infected crop (Skelsey et al., 2010) and a mutation rate of 1:10
9 , mutations
in any given gene of P. infestans are likely to occur in a disease-infected fi eld. Sexual
recombination, combined with the redundancy of several effectors that are recognized by
R -genes, explains why P. infestans easily mutates effector genes and develops virulence
(Jiang and Tyler, 2012).
As a result of the high genetic variation in potato, dozens of resistance sources have
been identifi ed and are available for breeding (Park et al., 2009). Whereas single genes
are easily defeated by virulent races to P. infestans , combinations of R -genes are more
effective, although the Pentland Dell case indicates that a more dynamic approach may
be needed (Niks et al., 2011).
Ideally, isogenic cultivars are developed that only differ in the combination of R -genes.
This would allow to deploy the most suitable cultivar, dependent on the epidemiology
of P. infestans . However, the introduction of one gene from a wild related species into a
tetraploid cultivar by traditional breeding already takes several decades, and to combine
different R -genes in a breeding programme is simply too complicated.
Since 1990, many R -genes to P. infestans have been mapped and cloned (Ballvora et al.,
2002; Huang et al., 2005; Park et al., 2009). These all belong to the so-called class of NBS/
LRR genes and have a cytoplasmic interaction with effector genes of P. infestans , resulting
in defence responses that block the growth of the pathogen (Jones and Dangl, 2006). A
genetic modifi cation (GM) approach to develop a series of isogenic cultivars with different
R -genes from crossable species is being pursued (Haverkort et al., 2016, Jacobsen and
Schouten, 2008). These so-called cisgenic plants might fall under the highly costly and
complex GM legislation, which would hamper their commercial opportunities.
The hybrid breeding system offers a clear path towards resilient resistance to P.
infestans : R -genes can be stacked in a potato hybrid via marker-assisted introgression
(Park et al., 2009). Two genes can be combined via the two parents in 2 3 years, and
additional R -genes can be added within a year to generate multi-stack resistance hybrids.
In this way, series of R -gene isogenic hybrids can be generated as a dynamic resource to
select the best combination of R -genes to protect the crop against the prevailing races of
P. infestans .
7.2 Hybrid potato breeding for East Africa
Hybrid potato cultivars will bring great benefi ts, not only to modern commercial farmers
in the developed world, but they may even have a greater social impact in tropical regions
where the population rely on potato as a major source of energy and nutrition (FAOstat,
2016). It is very challenging to start an initiative to develop hybrid potato for these regions
(Thomas-Sharma et al., 2016; Kumara et al., 2015). When legal and physical protection of
the breeding germplasm is not secured in these regions, the development of inbred lines
and the hybrid crosses are done elsewhere. The implementation of hybrid potato cultivars
in these regions requires considerable investments and strong cooperation of committed
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Hybrid potato breeding for improved varieties 17
public and private partners. As an example, the implementation of hybrid potato cultivars
for East Africa is briefl y described below.
The highlands of East Africa are traditionally important production areas for potato
because the crop is an important component of the local diet (Table 2). However, yields
remain very low (Table 2). A range of traditional varieties is used from local sources
(Kaguongo et al., 2008) as well as improved material from the International Centre for
Potato (CIP). Seed tubers are produced by farmers and storage conditions are far from
optimal (Kaguongo et al., 2008; Gildemacher et al., 2009). Janssens et al. (2013) concluded
that bacterial wilt, lack of clean seed tubers and poor storage are the most prominent
production constraints. Gildemacher et al. (2009) showed that only 3% of the seed tubers
sold were free of viruses.
True hybrid potato seeds are devoid of contaminating pathogens and therefore offer
an excellent opportunity to potato production improvement in East Africa. Such hybrids
should be attuned to the needs of the farmers, who grow their crop at the typical local
conditions like a short growing cycle of 90 100 days, high temperatures and tuber
development under short days. The prerequisites for establishing dedicated hybrid potato
breeding system for East Africa are:
A (private) organization executing a breeding programme tailored to the needs of
the region.
Secured supply of hybrid seeds for the region.
Regulations supporting imports and exports of seeds, seed tubers and commercial
Formal registration system for breeders rights protection.
Local organizations testing new experimental hybrids.
Such breeding programme can only become sustainable if the complete downstream part
of potato food chain is also well organized. This includes the following:
Production systems of disease-free seed tubers from seedlings.
An effi cient supply system for farmers to obtain hybrid cultivars.
Effi cient farmers cropping systems to produce high quality potato tubers for the
target markets.
Effi cient logistics to transport farm produce to consumers and processors.
Table 2 Potato area and production in six East-African countries in 2014
(FAOstat, 2016)
Country Area (000 ha) Production (000 tonnes) Yield (tonnes/ha)
Burundi 24.4 181.2 7.4
Kenya 115.6 1626.0 14.1
Rwanda 166.4 2225.1 13.4
Tanzania 211.5 1761.0 8.3
Uganda 39.0 188.0 4.8
Ethiopia 67.4 921.8 13.7
AQ: We
changed ‘T’
to ‘tonnes’
in “Table
2”. Please
check if this
is correct?
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18 Hybrid potato breeding for improved varieties
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A well-developed consumer market.
A business plan over the whole potato chain, whereby all stakeholders benefi t.
Solynta is already testing the fi rst experimental hybrids in the highlands of the Albertine
Rift in Ituri Province, Democratic Republic of Congo. These are experimental hybrids
derived from the European hybrid breeding programme. At a local farm, established by
the Lake Albert Foundation, seeds have been sown and seedlings have been transplanted
into the fi eld (Fig. 9). These hybrids are evaluated in good cooperation with local farmers
and the results are shared with the Solynta breeding programme. These data are used
to adjust the selection of inbred lines to the needs in East Africa and to continuously
generate new experimental hybrids, which will be tested at the Congo farm again. This
iterative and interactive process may already select the fi rst dedicated hybrids for East
Africa in 2 3 years.
This breeding programme is accompanied by research on cropping systems for the
region. In addition, training programmes for research institutions, agronomists, local staff
and interested farmers in the regions will make the farmers communities and relevant
institutions familiar with the new concepts of hybrid potato cultivars. Also market studies
are needed to identify and secure stable and sustainable markets for the farmers potato
products. This market may comprise other countries in the Great Lakes Region (South
Sudan, Uganda, Burundi and Rwanda).
The support for this programme by national authorities and development agencies
also in the neighbouring countries like Uganda is required, but it will take time before
the concept of hybrid potato cultivars is fully understood and accepted.
8 Conclusion
Since the fi rst crosses in 2008, the Solynta research efforts have been focused on the
development of a hybrid potato breeding system. This research has now reached the stage
where hybrid potato breeding will become reality. This has recently been supported by
two leading potato breeding companies in EU, KWS and HZPC, who have also expressed
their conviction that hybrid potato breeding will be future main breeding system (KWS,
2016). In addition, 21 leaders in the industry and potato science in the United States have
expressed their opinion on Reinventing potato as a diploid inbred line-based crop with a
scientifi c base for diploid hybrid potato breeding (Jansky et al., 2016).
We have made great advances in the development of useful homozygous inbred lines
and the fi rst fi eld evaluations of experimental hybrids have shown the potential of hybrid
cultivars to harvest heterosis.
The technologies to develop new cropping systems adapted to various climate zones
and agronomic practices, which allow the production of commercial seed tubers from
seedlings, are already available.
We envision that future hybrid potato varieties, similar to modern tomato hybrids, will
harbour up to 15 resistance genes. In addition, the lifetime of new potato hybrid cultivars
will be reduced to less than 5 years as is the experience in sugar beet, where the lifetime
of new cultivars is only 2 3 years since the fi rst diploid hybrids have been introduced into
the market.
Our inbred lines will also be of great value for research purposes as they allow the
development of sophisticated populations that are very helpful in genetic and genomic
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Hybrid potato breeding for improved varieties 19
studies as has been shown in other crops. This will give a boost in the exploration and
exploitation of the genetic variation in the potato germplasm.
The self-compatible potato inbred lines will greatly stimulate quantitative research on
the genetic control of complex traits such as interaction with mycorrhiza, biotic stresses,
nutrient uptake, nutrition value and processing quality.
The application of hybrid potato breeding technology will not be restricted to the most
advanced research groups in the public or private institutions. New players in the scientifi c
and applied breeding fi eld of the potato business will arise and new cooperations will
be established to take full advantage of the hybrid breeding technology in science and
product development.
Finally, hybrid potato breeding will require the skills of the breeder as well as the
knowledge and tools of scientists. As a result, a new generation of potato breeding teams
will be established combining the skills of the breeder with the in-depth knowledge of
plant scientists.
9 Where to look for further information
This chapter describes the fi rst implementation of a hybrid potato breeding strategy
and the progress that is made since 2008. We direct the interested reader to the papers
of Almekinders (1996, 2009) to learn about conventional TPS . As a textbook on plant
breeding, we suggest Brown and Cagliari (2011), while more advanced potato genetics
and genomics is found in Bradeen and Kole (2011). The history and mechanism of
Phytophthora attack is well described by Fry (2008). Finally, FAO (FAOstat, 2016) statistics
provide numerous data on potato cultivation.
Leading centres of research include Wageningen University in The Netherlands, James
Hutton Institute in the United Kingdom, the International Potato Centre in Peru, Wisconsin
University and Michigan State University both in the United States, whereby the most
recent paper of Jansky et al. (2016) can be considered a must for anybody interested in
hybrid potato breeding.
10 Acknowledgements
We are grateful to Jan Leemans and Herman Fleer for critically reading and reviewing this
11 References
Abrol, D. P. (2012). Pollination for hybrid seed production. In Pollination Biology. Biodiversity
Conservation and Agricultural Production . Publisher Springer Netherlands, pp. 397 411.
Almekinders, C. J. M., Chujoy, E. and Thiele, G. (2009). The use of true potato seed as pro-poor
technology: The efforts of an international agricultural research institute to innovating potato
production. Pot. Res. 52, 275 93.
Almekinders, C. J. M., Chilver, A. S. and Renia, H. M. (1996). Current status of the TPS technology in
the world. Pot. Res. 39, 289 303.
Chapter 4_potatoes vol 1.indd 19Chapter 4_potatoes vol 1.indd 19 26-09-2016 12:47:1126-09-2016 12:47:11
20 Hybrid potato breeding for improved varieties
© Burleigh Dodds Science Publishing Limited, 2016. All rights reserved.
Amin, N., Amin, A. R., Roy, T. S., Ali, M. A., Rashid, M. M., Hossain, M. M. and Hasan, N. (2014).
Bulking behavior of seedling tubers derived from true potato seed as affected by its size and
harvesting time. App. Sci. Rep. 8, 1 8.
Ballvora, A., Ercolano, M. R., Weiss, J., Meksem, K., Bormann, C. A., Oberhagemann, P., Salamini, F.
and Gebhardt, C. (2002). The R1 gene for potato resistance to late blight ( Phytophthora infestans )
belongs to the leucine zipper/NBS/LRR class of plant resistance genes. Plant J. 30, 361 71.
Belhaj, K., Chaparro-Garcia, A., Kamoun, S. and Nekrasov, V. (2013). Plant genome editing made
easy: targeted mutagenesis in model and crop plants using the CRISPR/Cas system. Plant Meth.
9, 39.
Birchler, J. A., Yao, H., Chudalayandi, S., Vaiman, D. and Veitia, R. A. (2010). Heterosis. Plant Cell 22,
2105 12.
Bradeen, J. M. and Kole, C. (2011). Potato Genetics: Genetics, Genomics and Breeding of Potato .
CRC Press, USA
Bradshaw, J. E., Hackett, C. A., Pande, B., Waugh, R. and Bryan, G. J. (2007). QTL mapping of yield,
agronomic and quality traits in tetraploid potato ( Solanum tuberosum subsp. tuberosum ). Theor.
Appl. Genet. 116, 193 211.
Brown, J. and Caligari, P. (2011). An Introduction to Plant Breeding . Wiley-Blackwell, USA, p. 224.
Cappadocia, M. (1990). Wild Potato ( Solanum chacoense Bitt.). In Y. P. S. Bajaj (ed.), Vitro Production
of Haploids. Biotechnology in Agriculture and Forestry, Vol. 12 Haploids in Crop Improvement I ,
pp. 514 29. Springer-Verlag, Berlin, Heidelberg.
Chani, E., Veilleux, R. E. and Boluarte-Medina, T. (2000). Improved androgenesis of interspecifi c
potato and effi ciency of SSR markers to identify homozygous regenerants. Plant Cell Tissue
Organ Cult. 60, 101 12.
Charlesworth, D. and Willis, J. H. (2009). The genetics of inbreeding depression. Genetics 10, 783 96.
Chauvin, J. E., Souchet, C., Dantec, J. P. and Ellisseche, D. (2003). Chromosome doubling of 2x
Solanum species by oryzalin: method development and comparison with spontaneous
chromosome doubling in vitro . Plant Cell Tissue Organ Cult. 73, 65 73.
Chen, X., Niks, R. E., Hedley, P. E., Morris, J., Druka, A., Marcel, T. C., Vels, A. and Wauh, R. (2010).
Differential gene expression in nearly isogenic lines with QTL for partial resistance to Puccinia
hordei in barley. Genomics 11, 629.
Crow, J. F. (1998). 90 Years ago: The beginning of hybrid maize. Genetics 148, 923 8.
CSAC (2005). Initial sequence of the chimpanzee genome and comparison with the human genome.
Nature 437, 69 87.
D hoop, B. B., Keizer, P. L. C., Jo ã o Paulo, M., Visser, R. G. F., Van Eeuwijk, F. A. and Van Eck, H. J.
(2014). Identifi cation of agronomically important QTL in tetraploid potato cultivars using a
marker-trait association analysis. Theor. Appl. Genet. 127, 731 48.
De Jong, H. and Rowe, P. R. (1971). Inbreeding in cultivated diploid potatoes. Pot. Res. 14, 74 83
De Mainea, M. J. (1982). An evaluation of the use of dihaploids and unreduced gametes in breeding
for quantitative resistance to potato pathogens. J. Agric. Sci. 99, 79 83.
Delleman, J., Mulder, A. and Turkensteen, L. J. (2004). Potato Diseases: Diseases, Pests and Defects .
Potatoworld and NIVAP, The Hague, the Netherlands.
Douches, D. S., Maas, D. J., Astrzebski, K. and Chase, R. W. (1996). Assessment of potato breeding
progress in the USA over the last century. Crop Sci. 36, 1544 52.
Duncan, J. M. (1999). Phytophthora -an abiding threat to our crops. Microbiol. Today 26, 114 16.
Duvick, D. N. (2005). The contribution of breeding to yield advances in maize (Zea mays L.). Adv.
Agron. 86, 83 145.
El-Bedewy, R., Crissman, C. and Cortbaoui, R. (1994). Progress report. Egypt s seed system based on
true potato seed. CIP Circular 20, 5 8.
El-Yazied, A., Elminiawy, S. E., Hamoud, N. K. and El-Kheima, S. (2004). Seed tuber production of
some hybrids using true potato seed. Mansoura University. J. Agric. Sci. 32, 1329 41.
Endelman, J. B. and Jansky, S. H. (2016). Genetic mapping with an inbred line-derived F
2 population
in potato. Theor. Appl. Genet. 1 9
Chapter 4_potatoes vol 1.indd 20Chapter 4_potatoes vol 1.indd 20 26-09-2016 12:47:1126-09-2016 12:47:11
© Burleigh Dodds Science Publishing Limited, 2016. All rights reserved.
Hybrid potato breeding for improved varieties 21
FAO, IFAD and WFP. (2015). The State of Food Insecurity in the World 2015. Meeting the 2015
international hunger targets: taking stock of uneven progress. Rome, FAO.
FAOstat. (2016).
Finkers, R., Van Heusden, A. W., Meijer-Dekens, F., Van Kan, J. A. L. and Lindhout, P. (2007). The
construction of a Solanum habrochaites LYC4 introgression population and the identifi cation of
QTLs for resistance to Botrytis cinerea. Theor. Appl. Genet. 114, 1071 80.
Frisch, M. and Melchinger, A. E. (2005). Selection theory for marker-assisted backcrossing. Genetics
170, 909 17.
Fry, W. (2008). Plant diseases that changed the world Phytophthora infestans : the plant (and R gene)
destroyer. Molec. Pl. Path. 9, 385 402.
Gildemacher, P. R., Demo, P., Barker, I., Kaguongo, W., Woldegiorgis, GT., Wagoire, W. W., Wakahiu, M.,
Leeuwis, C. and Struik, P. C. (2009). A Description of seed potato systems in Kenya, Uganda and
Ethiopia. Am. J. Pot. Res. 86, 373 82.
Gopal, J. (2004). True potato seed: Breeding for hardiness. In A. Haneafi (ed.), Sixth Triennial Congress
of the African Potato Association. Proc. APA Congr. , 5 10 April, Agadir, Morocco, pp. 39 57.
Gopal, J. (2014). Heterosis breeding in potato. Agric. Res. 3, 204 17.
Haldane, J. (1930). Theoretical genetics of autopolyploids. J. Genet. 22, 359 72.
Haverkort, A. J., Boonekamp, P. M., Hutten, R., Jacobsen, E., Lotz, L. A. P., Kessel, G. J. T., Vossen,
J. H. and Visser, R. G. F. (2016). Durable late blight resistance in potato through dynamic
varieties obtained by cisgenesis: Scientifi c and societal advances in the DuRPh project. Pot.
Res. , 59, 35 66.
Hawkes, J. G. (1990). The Potato: Evolution, Biodiversity, and Genetic Resources . Belhaven Press,
Hosaka, K. and Hanneman, R. E. (1998a). Genetics of self-compatibility in a self-incompatible wild
diploid potato species Solanum chacoense . 1. Detection of an S locus inhibitor ( Sli ) gene.
Euphytica 99, 191 7.
Hosaka, K. and Hanneman, R. E. (1998b). Genetics of self-compatibility in a self-incompatible wild
diploid potato species Solanum chacoense . 2. Localization of an S locus inhibitor ( Sli ) gene on
the potato genome using DNA markers. Euphytica 103, 265 71.
Hua, J., Xing, Y., Wu, W., Xu, C., Sun, X., Yu, S. and Zhang, Q. (2003). Single-locus heterotic effects
and dominance by dominance interactions can adequately explain the genetic basis of heterosis
in an elite rice hybrid. Proc. Natl. Acad. Sci. USA 100, 2574 9.
Huang, S., Van der Vossen, E. A., Kuang, H., Vleeshouwers, V. G., Zhang, N., Borm, T. J., Van Eck, H. J.,
Baker, B., Jacobsen, E. and Visser, R. G. (2005). Comparative genomics enabled the isolation of
the R3a late blight resistance gene in potato. Plant J. 42, 251 61.
Hutten, R. C. B. (1994). Basic Aspects of Potato Breeding Via the Diploid Level. PhD Thesis,
Wageningen University, p. 93.
Hutten, R. C. B., Schippers, M. G. M., Hermsen, J. G. Th . and Jacobsen, E. (1994). Comparative
performance of diploid and tetraploid progenies from 2x.2x crosses in potato. Euphytica 81,
187 92.
Jacobsen, E. and Schouten, H. J. (2008). Cisgenesis, a new tool for traditional plant breeding, should
be exempted from the regulation on genetically modifi ed organisms in a step by step approach.
Pot. Res. 51, 75.
Jansky, S. H. and Peloquin, S. J. (2006). Advantages of wild diploid Solanum species over cultivated
diploid relatives in potato breeding programs. Genet. Res. Crop Evol. 53, 669 74.
Jansky, S. H., Charkowski, A. O., Douches, D. S., Gusmini, G., Richael, C., Bethke, P. C., Spooner,
D. M., Novy, R. G., De Jong, H., De Jong, W. S., Bamberg, J. B., Thompson, A. L., Bizimungu,
B., Holm, D. G., Brown, C. R., Haynes, K. G., Sathuvalli, V. R., Veilleux, R. E., Miller Jr., J. C.,
Bradeen, J. M. and Jiang, J. M. (2016). Reinventing potato as a diploid inbred line-based crop.
Crop Sci. 56, 1 11.
Jansky, S. H., Chung, Y. S. and Kittipadakul, P. (2014). M6: A diploid potato inbred line for use in
breeding and genetics research. J. Plant Registr. 8, 195 9.
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22 Hybrid potato breeding for improved varieties
© Burleigh Dodds Science Publishing Limited, 2016. All rights reserved.
Janssens, S. R. M., Wiersema, S. G., Goos, H. and Wiersema, W. (2013). The value chain for seed
and ware potatoes in Kenya; Opportunities for development LEI. Memorandum 13-080, p. 57.
Jeong, H-S., Jang, S., Han, K., Kwon, J-K. and Kang, B-C. (2015). Marker-assisted backcross breeding
for development of pepper varieties ( Capsicum annuum ) containing capsinoids. Molec.
Breeding 35, 226.
Jeuken, M. J. W. and Lindhout, P. (2004). The development of lettuce backcross inbred lines (BILs) for
exploitation of the Lactuca saligna (wild lettuce) germplasm. Theor. Appl. Genet. 109, 394 401.
Jiang, R. H. Y. and Tyler, B. M. (2012). Mechanisms and evolution of virulence in oomycetes. Ann. Rev.
Phytop. 50, 295 318.
Jones, J. D. G and Dangl, J. L. (2006). The plant immune system. Nature 444, 323 9.
Kaguongo, W., Gildemacher, P., Demo, P., Wagoire, W., Kinyae, P., Andrade, J., Forbes, G., Fuglie, K., and
Thiele, G. (2008). Farmer practices and adoption of improved potato varieties in Kenya and Uganda.
International Potato Center (CIP), Lima, Peru. Social Sciences Working Paper 2008 5. 85 p.
Kumar, V. (2014). True potato seed technology Prospects and problems. In N. K. Pandey, D. K. Singh
and R. Kumar (eds), Current Trends in Quality Potato Production, Processing and Marketing ,
pp. 175 82. Central Potato Research Institute, India.
Kumara, N. S., Govindakrishnan, P. M., Swarooparani, D. N., Nitin, Ch. Surabhi, J. and Aggarwal, P. K.
(2015). Assessment of impact of climate change on potato and potential adaptation gains in the
Indo-Gangetic Plains of India. Intern. J. Pl. Prod. 9 (1), 151–70.
KWS press release (2016).
Leskovar, I. D., Crosby, M. K., Palma, A. M. and Edelstein, M. (2014). Vegetable crops: Linking
production, breeding and marketing. In R. G. Dixon and E. D. Aldous (eds), Horticulture: Plants
for people and places, Volume 1: Production Horticulture , pp. 75 96. Springer Netherlands,
Li, L., Paulo, M.-J., Van Eeuwijk, F., and Gebhardt, C. (2010). Statistical epistasis between candidate
gene alleles for complex tuber traits in an association mapping population of tetraploid potato.
Theor. Appl. Genet. 121, 1303 10.
Li, X.-Q. (2008). Male sterility systems for hybrid seed production in Brassica crops. CAB Reviews:
Perspectives in Agriculture, Veterinary Science, Nutrition and Natural Resources 3, 1 14.
Lindhout, P., Meijer, D., Schotte, T., Hutten, R. C. B., Visser, R. G. F. and Van Eck, H. J. (2011a). Towards
F1 hybrid seed potato breeding. Pot. Res. 54, 301 12.
Lindhout, W. H., Schotte, T. P., Visser, R. G. F., Van Eck, H. J. and Hutten, R. C. B. (2011b). Hybrid seed
potato breeding. European Patent Offi ce nr EP 2514303 A1
Lippman, Z. B. and Zamir, D. (2007). Heterosis: revisiting the magic. Trends Genet. 23, 60 6.
Lommen, W. J. M. (1995). Basic studies on the production and performance of potato minitubers.
PhD Thesis, Wageningen Agricultural University, Wageningen, The Netherlands, p. 181.
Lommen, W. J. M. (2007). The canon of potato science: Hydroponics. Pot. Res. 50, 315 18.
L ö ssl, A., G ö tz, M., Braun, A. and Wenzel, G. (2000). Molecular markers for cytoplasm in potato: Male
sterility and contribution of different plastid-mitochondrial confi gurations to starch production.
Euphytica 116, 221 30.
Malcolmson, J. F. (1969). Races of Phytophthora infestans occurring in Great Britain. Trans. Br. Mycol.
Soc. 53, 417 23.
Mallick, N., Vinod, Sharma, J. B., Tomar, R. S., Sivasamy, M. and Prabhu, K. V. (2015). Marker-assisted
backcross breeding to combine multiple rust resistance in wheat. Plant Breed. 134, 172 7.
Meijer, D. A., Abdullah, S., Rothengatter, R., Van Eck, H. J., Visser, R. G. F., Lindhout, P. and Van
Heusden, S. (2016). An F
2 QTL mapping study in a cross between S. tuberosum and S.
chacoense . Theor. Appl. Genet. (submitted).
Muthoni, J., Shimelis, H., Melis, R. and Kinyua, Z. M. (2014). Response of potato genotypes to bacterial
wilt caused by Ralstonia solanacearum (Smith) (Yabuuchi et al.) in the tropical highlands. Am. J.
Pot. Res. 91, 215 32.
Chapter 4_potatoes vol 1.indd 22Chapter 4_potatoes vol 1.indd 22 26-09-2016 12:47:1126-09-2016 12:47:11
© Burleigh Dodds Science Publishing Limited, 2016. All rights reserved.
Hybrid potato breeding for improved varieties 23
Niks, R. E., Parlevliet, J. E., Lindhout, P. and Bai, Y. (2011). Breeding Crops with Resistance to Diseases
and Pests . Wageningen Academic Publishers, p. 200.
Paran, I., Goldman, I., Tanksley, S. D. and Zamir, D. (1995). Recombinant inbred lines for genetic
mapping in tomato. Theor. Appl. Genet. 90, 542 8.
T. H., Vleeshouwers, V. G. A. A., Jacobsen, E., Van der Vossen, E. and Visser, R. G. F. (2009).
Molecular breeding for resistance to Phytophthora infestans (Mont.) de Bary in potato ( Solanum
tuberosum L.): a perspective of cisgenesis. Plant Breed. 128, 109 17.
PGSC (2011). Genome sequence and analysis of the tuber crop potato. Nature 475, 189 94.
Phumichai, C. and Hosaka, K. (2006). Cryptic improvement for fertility by continuous selfi ng of diploid
potatoes using Sli gene. Euphytica 149, 251 8.
Phumichai, C., Mori, M., Kobayashi, A., Kamijima, O. and Hosaka, K. (2005). Toward the development
of highly homozygous diploid potato lines using the self-compatibility controlling Sli gene.
Genome 48, 977 84.
Prashar, A., Hornyik, C., Young, V., McLean. K., Kumar Sharma, S., Dale, M. F. B. and Bryan, G. J.
(2014). Construction of a dense SNP map of a highly heterozygous diploid potato population
and QTL analysis of tuber shape and eye depth. Theor. Appl. Genet. 127, 2159 71.
Renia, H. (1995). True seed is a commercial reality in USA. Pot. Rev. 5, 48 51.
Rijk, B., van Ittersum, M. and Withagen, J. (2013). Genetic progress in Dutch crop yields. Field Crops
Res. 149, 262 8.
Rowe, P. R. (1967). Performance and variability of diploid and tetraploid potato families. Am. Pot. J.
44, 263 71.
Roy, T. S., Baque, M. A., Chakraborty, R., Haque, M. N. and Suter, P. (2015). Yield and economic return
of seedling tuber derived from True Potato Seed as infl uenced by tuber size and plant spacing.
Univ. J. Agric. Res. 3, 23 30.
Schmalenbach, I. and Pillen, K. (2009). Detection and verifi cation of malting quality QTLs using wild
barley introgression lines. Theor. Appl. Genet. 118, 1411 27.
Skelsey, P., Rossing, W. A. H., Kessel, G. J. T. and Van der Werf, W. (2010). Invasion of Phytophthora
infestans at the landscape level: How do spatial scale and weather modulate the consequences
of spatial heterogeneity in host resistance? Phytopath. 100, 1146 61.
Solomon-Blackburn, R. M. and Barker, H. (2001). A review of host major-gene resistance to potato
viruses X, Y, A and V in potato: genes, genetics and mapped locations. Heredity 86, 8 16.
Struik, P. C. and Wiersema, S. (2012). Seed Potato Technology . Wageningen Academic Publishers,
p. 383.
Su, C. F., Lu, W. G., Zhao, T. J. and Gai, J. Y. (2009). Verifi cation and fi ne-mapping of QTLs conferring
days to fl owering in soybean using residual heterozygous lines. Chin. Sci. Bull. 6, 499 508.
Thomas-Sharma, S., Abdurahman, A., Ali, S., Andrade-Piedra, J. L., Bao, S., Charkowski, A. O.,
Crook, D., Kadian, M., Kromann, P., Struik, P. C., Torrance, L., Garrett, K. A. and Forbes, G. A.
(2016). Seed degeneration in potato: the need for an integrated seed health strategy to mitigate
the problem in developing countries. Plant Path. 65, 3 16.
Tierno, R., Carrasco, A., Ritter, E. and Ruiz de Galarreta, J. I. (2014). Differential growth response and
minituber production of three potato cultivars under aeroponics and greenhouse bed culture.
Amer. J. Pot. Res. 91, 346 53.
Tobias, A., Schrag, T. A., M ö hring, J., Melchinger, A. E., Kusterer, B., Dhillon, B. S., Piepho, H-P. and
Frisch, M. (2009). Prediction of hybrid performance in maize using molecular markers and joint
analyses of hybrids and parental inbreds. Theor. Appl. Genet. 120, 451 61.
Troyer, A. F. (2006). Adaptedness and heterosis in corn and mule hybrids. Crop Sci . 46, 528 43.
Uijtewaal, B. A., Huigen, D. J. and Hermsen, J. G. Th. (1987a). Production of potato monohaploids
(2n = x = 12) through prickle pollination. Theor. Appl. Genet. 73, 751 8.
Uijtewaal, B. A., Jacobsen, E. and Hermsen, J. G. Th (1987b). Morphology and vigour of monohaploid
potato clones, their corresponding homozygous diploids and tetraploids and their heterozygous
diploid parent. Euphytica 36, 745 53.
Chapter 4_potatoes vol 1.indd 23Chapter 4_potatoes vol 1.indd 23 26-09-2016 12:47:1126-09-2016 12:47:11
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© Burleigh Dodds Science Publishing Limited, 2016. All rights reserved.
Uitdewilligen, J. G. A. M. L., Wolters, A. M. A, D hoop, B. B., Borm, T. J. A., Visser, R. G. F. and
Van Eck, H. J. (2013). A next-generation sequencing method for genotyping-by-sequencing of
highly heterozygous autotetraploid potato. PLoS ONE 8(5), e62355.
Van Breukelen, E. W. M., Ramanna, M. S. and Hermsen, J. G. Th . (1977). Pathenogenetic monohaploids
(2n = 2x = 12 from Solanum tuberosum L. and S. verrucosum Schlechtd. and the production of
homozygous potato diploids. Euphytica 26, 263 71.
Van Eck, H. J., Jacobs, J. M. E., van den Berg, P. M. M. M., Stiekema, W. J. and Jacobsen, E. (1994),
The inheritance of anthocyanin pigmentation in potato ( Solanum tuberosum L.) and mapping of
tuber skin colour loci using RFLPs. Heredity 73, 410 21.
Van Eck, H. J., Jacobs, J. M. E., van Dijk, J., Stiekema, W. J. and Jacobsen, E. (1993) Identifi cation and
mapping of three fl ower colour loci of potato ( S. tuberosum L.) by RFLP analysis. Theor. Appl.
Genet. 86, 295 300.
V í quez-Zamora, M., Caro, M., Finkers, R., Tikunov, Y., Bovy, A., Visser, R. G. F., Bai, Y. and Van
Heusden, S. (2014). Mapping in the era of sequencing: high density genotyping and its application
for mapping TYLCV resistance in Solanum pimpinellifolium . BMC Genomics 15, 1152.
Visser, R. G. F., Bachem, C. W. B., Borm, T., de Boer, J., Van Eck, H. J., Finkers, R., Van der Linden, G.,
Maliepaard, C. A., Uitdewilligen, J. G. A. M. L., Voorrips, R., Vos, P. and Wolters, A. M. A. (2014).
Possibilities and challenges of the potato genome sequence. Pot. Res. 57, 327 30.
Vos, P. G., Uitdewilligen, J. G. A. M. L, Voorrips, R. E., Visser, R. G. F. and Van Eck, H. J. (2015).
Development and analysis of a 20K SNP array for potato ( Solanum tuberosum ): an insight into
the breeding history. Theor. Appl. Genet. 128, 2387 401.
Wang, C. M., Lo, L. C, Feng, F., Zhu, Z. Y. and Yue, G. H. (2008). Identifi cation and verifi cation of
QTL associated with growth traits in two genetic backgrounds of Barramundi ( Lates calcarifer ).
Animal Genet. Vol. 39, 34 9.
Wang om, W. G. and van Dijk, M. P. (2013). Low potato yields in Kenya: do conventional input
innovations account for the yields disparity? Agric. Food Sec. 2, 14.
Wiersema, S. G. (1986). A method of producing seed tubers from true potato seed. Pot. Res. 29,
225 37.
Young, N. D., Zamir, D., Ganal, M. W. and Tanksley, S. D. (1988). Use of isogenic lines and simultaneous
probing to identify DNA markers tightly linked to the tm-2a gene in tomato. Genetics 120,
579 85.
Zhang, Y-M., Mao, Y., Xie, C., Smith, H., Luo, L. and Xu, S (2005). Mapping Quantitative Trait Loci
using naturally occurring genetic variance among commercial inbred lines of maize (Zea mays
L.). Genetics 169, 2267 75.
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... To render this germplasm self-compatible, it was crossed with a wild potato species carrying a self-compatibility restorer gene Sli, which was made available by the Japanese researchers who identified this gene [1]. Thus, the company was able to initiate a hybrid potato breeding program by generating diploid homozygous inbred lines [4]. With the development of a sufficient number of highly inbred lines, it became possible to systematically combine genes and exploit heterosis in hybrid varieties, also introducing desirable traits such as disease resistance by marker-assisted introgression breeding [12][13][14]. ...
... The start of a hybrid breeding program based on self-compatible diploid potatoes is described both by outside observers and the Solynta team itself as a revolutionary 'paradigm shift', involving the transformation of breeding, seed and cultivation as three interrelated systems [4,5,30]. Firstly, it redefines the traditional and time-consuming tetraploid potato breeding system. ...
... Key steps in hybrid potato development have also been enabled by skills, knowledge and (gene) technologies that are widely available, for example, molecular DNA 'markers'. In the context of hybrid breeding, such markers can be used to test the level of homozygosity in inbred parental lines and also to predict more accurately the breeding value of these lines [4]. Moreover, by so-called marker-assisted introgression breeding, publicly available favorable traits can be introduced into these lines, for example, the Phytophthora resistance genes that were stacked by Solynta in their potato parental lines [12]. ...
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Citation: Stemerding, D.; Beumer, K.; Edelenbosch, R.; Swart, J.A.A.; de Vries, M.E.; ter Steeg, E.; Almekinders, C.J.M.; Lindhout, P.; van Dijk, L.C.M.; Struik, P.C. Responsible Innovation in Plant Breeding: The Case of Hybrid Potato Breeding. Plants 2023, 12, 1751. Abstract: As an emerging innovation, hybrid potato breeding raises high expectations about faster variety development and clean true potato seed as a new source of planting material. Hybrid breeding could, therefore, substantially contribute to global food security and other major sustainable development goals. However, its success will not only depend on the performance of hybrid potato in the field, but also on a range of complex and dynamic system conditions. This article is based on a multidisciplinary project in which we have studied the innovation dynamics of hybrid potato breeding and explored how these dynamics may shape the future of hybrid potato. Inspired by the approach of responsible innovation, we closely involved key players in the Dutch and international potato sector and other relevant actors in thinking about these potato futures. An important and recurrent theme in our work is the tension between the predominant commercial innovation dynamics in plant breeding and promises to respond to the global challenges of food security, agrobiodiversity and climate change. In this article, we, therefore, discuss responsible innovation strategies in (hybrid) potato breeding, which may help to bridge this tension and finally reflect on the implications for the field of plant breeding in general.
... For the introduction of HTPS, an accessible and reliable seed system is pivotal to provide farmers with the desired starting material, which could be clean seeds, seedlings or seedling tubers (Lindhout et al., 2018). Any intervention involving seeds or starting material should build on a solid understanding of existing seed systems . ...
... Update variety selection criteria sustainable introduction and continued use of HTPS (Lindhout et al., 2018). This poses major challenges because currently, commercialisation of HTPS is in its infancy. ...
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Hybrid true potato seed (HTPS) holds the promise to foster development in potato sectors in East-Africa. Compared to a conventional production system based on clonal propagation of tetraploid varieties, clear advantages of HTPS are easier transport and storage of starting material, high multiplication rates, increased availability of disease-free planting material and faster and more targeted breeding. Despite these advantages, an informed assessment of how HTPS can be implemented into East African seed and farming systems, and of what the enabling conditions are for such implementation is still necessary. Our aim is to understand if and how HTPS could foster development of potato sectors in East Africa and what are the major opportunities and bottlenecks. We describe the context of potato production in East Africa, and we summarise the key lessons from past experiences with the implementation of (non-hybrid) true potato seed. This informs our analysis of the requirements for an effective and inclusive introduction. Major requirements are a solid understanding of: (1) the criteria farmers use to assess the innovation; (2) local farm realties; and (3) the seed system in which farmers operate. The implementation of HTPS requires adaptations in agronomic management and in seed system configurations, but these adaptations need to fit into the agro-ecological and socio-economic context. This can be achieved through an iterative research cycle linking farmers, researchers, breeders and other stakeholders. To facilitate institutional embedding of the innovation, interactive, multi-stakeholder processes are required to develop shared views on the acceptability, sustainability and societal desirability of HTPS. We conclude by drawing a research agenda with urgent questions that need empirical research prior to and during the introduction of HTPS in East Africa.
... Impact of hybrid potato which makes available potato starting material for cultivation in the form of diploid hybrid true potato seed (Lindhout et al., 2011;Jansky et al., 2016;Lindhout et al., 2018). This seed is devoid of (almost all) contaminating pathogens and pests and can be rapidly propagated, and easily transported and stored. ...
... This seed is devoid of (almost all) contaminating pathogens and pests and can be rapidly propagated, and easily transported and stored. The diploid hybrid potato also promises to significantly accelerate the process of breeding compared to conventional practices today (Lindhout et al., 2018;Su et al., 2020). As a result, this new hybrid potato may provide a highly promising response to the global challenges of food security, sustainability and climate change. ...
... Both trials were performed with four diploid test-hybrids (H1, H2, H3, H4) that were produced in a diploid hybrid breeding program as described by Lindhout et al. (2011Lindhout et al. ( , 2018. All hybrids produce medium -high yields. ...
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For potato, diploid hybrid breeding is a novel breeding technique that speeds up the development of new varieties. A consequence of hybrid breeding is the introduction of hybrid true potato seeds as starting material. From these seeds, seedling tubers can be produced in one field season, to use as starting material for a seed or a ware crop in the following year. For breeding purposes as well as for seed crop and ware crop production it is essential to produce seedling tubers of high quality. The production of seedling tubers is a new step in the potato production chain, therefore, we investigated the effect of tuber quality traits on plant development and yield. With similar seedling tuber weight, more eyes per seedling tuber led to more stems per plant. This was compensated by a lower number of tubers per stem resulting in an equal total tuber number and weight per plant at the end of the growing season. A higher seedling tuber weight led to a higher soil cover in the field. Therefore, all hybrids produced a higher total tuber weight per plant when grown from larger seedling tubers than when grown from smaller seedling tubers, while number of eyes and stems per tuber has no effect on final yield when using equal seedling tuber weight.
... Breeding programs considering changing to hybrid strategies to increase genetic gain per unit cost need additional research to inform their decisions. This is especially true for clonal crops, which show inbreeding depression and heterosis in some economically important traits but have unique features that may decrease the competitiveness of hybrid breeding strategies such as autopolyploidy, delayed flowering, and low multiplication rates (Aighewi et al. 2015;Diaz et al. 2021;Ceballos et al. 2015Ceballos et al. , 2020Darkwa et al. 2020;Batte et al 2020;Lindhout et al. 2018;Werner et al. 2023;Amadeu et al. 2020). ...
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Key message Reciprocal recurrent selection sometimes increases genetic gain per unit cost in clonal diploids with heterosis due to dominance, but it typically does not benefit autopolyploids. Abstract Breeding can change the dominance as well as additive genetic value of populations, thus utilizing heterosis. A common hybrid breeding strategy is reciprocal recurrent selection (RRS), in which parents of hybrids are typically recycled within pools based on general combining ability. However, the relative performances of RRS and other breeding strategies have not been thoroughly compared. RRS can have relatively increased costs and longer cycle lengths, but these are sometimes outweighed by its ability to harness heterosis due to dominance. Here, we used stochastic simulation to compare genetic gain per unit cost of RRS, terminal crossing, recurrent selection on breeding value, and recurrent selection on cross performance considering different amounts of population heterosis due to dominance, relative cycle lengths, time horizons, estimation methods, selection intensities, and ploidy levels. In diploids with phenotypic selection at high intensity, whether RRS was the optimal breeding strategy depended on the initial population heterosis. However, in diploids with rapid-cycling genomic selection at high intensity, RRS was the optimal breeding strategy after 50 years over almost all amounts of initial population heterosis under the study assumptions. Diploid RRS required more population heterosis to outperform other strategies as its relative cycle length increased and as selection intensity and time horizon decreased. The optimal strategy depended on selection intensity, a proxy for inbreeding rate. Use of diploid fully inbred parents vs. outbred parents with RRS typically did not affect genetic gain. In autopolyploids, RRS typically did not outperform one-pool strategies regardless of the initial population heterosis.
... In all trials, inbred lines (IL1-IL11) were used that were obtained after several generations of self-pollination by Solynta, as described by Lindhout et al. (2011Lindhout et al. ( , 2018. All lines were developed by manual selfpollinating for between 6 and 10 generations. ...
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Growing small seedling tubers from true seed, comparable with mini tubers, in controlled conditions could be a method to multiply healthy starting material of potato. In indoor farming systems, the conditions can be optimized for high production. In field trials it is impossible to investigate the effects of environmental factors such as temperature and light separately. In this study, we performed three climate room experiments in which the effects of light intensity, temperature and percentage of far-red light in the light spectrum on tuber production were assessed. We found that increasing the average temperature reduced tuber number and tuber weight. Increasing the diurnal temperature variation while keeping the average temperature equal resulted in increased tuber size. The light treatments on the other hand only affected the number of tubers per plant: increasing light intensity and increasing the percentage of far-red light in the spectrum enhanced the number of tubers. Moreover, interaction in tuber production between inbred lines and temperature was significant, with some inbred lines being relatively tolerant to high temperature. These findings will help breed for heat tolerant varieties and optimize growing conditions for tuber production in indoor farming systems.
... The main disadvantage of OP TPS was the high heterogeneity of the material, especially with the increased need for highly uniform cultivars in the processing markets . However, these challenges are now addressed through the introduction of hybrid diploid breeding (Lindhout et al., 2011), through which the resulting F1 hybrid TPSs are highly uniform and still maintain the many advantages of OP TPS (see Lindhout et al., 2018). Results of the studies on the feasibility of the TPS-based systems highlighted multiple advantages of the individual cultivation pathways and the challenges that should be addressed to ensure the success of the systems (Almekinders et al., 1996;Struik & Wiersema, 1999;Almekinders et al., 2009). ...
Full-text available
The introduction of hybrid breeding in potato (Solanum tuberosum L.) requires novel and efficient cropping systems for potato production based on true potato seed (TPS). Such systems address the limitations of conventional seed-tuber based systems including low multiplication rates, high degeneration rates and high costs of transport and storage. Of the possible cultivation pathways of TPS, we introduce and discuss the potential of field transplanting nursery-raised potato seedlings as an alternative system for seed and ware production. This review discusses the current knowledge available on field transplanting of potato seedlings, the key factors that influence the success or failure of the system, and some of the prospective factors that will influence the wide introduction and utilization of field transplanting of potato seedlings in diverse farming systems. A field transplanting system will require successful production of seedlings in the nursery, successful establishment of transplanted seedlings in the field and successful crop management to attain a productive seed or ware crop. The contribution of various factors in the various phases of the system to the success of the transplanted crop is also discussed. The introduction and utilization of the field transplanting system will be accelerated when hybrid breeding focuses on introgression of traits of interest into high yielding cultivars and when agronomic studies focus on defining factors influencing productivity in distinct phases of the system.
... This different type of starting material further complicates the application of 18 Impact of hybrid potato hybrid cultivars in potato tuber production, although it strongly increases the multiplication rate, thereby reducing the number of years between making the cross and producing enough seed for commercial use. The limitations caused by tetrasomic inheritance and inbreeding depression have been overcome and at present, there are hybrid potato breeding programmes in the Netherlands, USA and China (Jansky et al., 2016;Lindhout et al., 2011Lindhout et al., , 2018Zhang et al., 2021). ...
... Thus, as a general conclusion, most of our interviewees stressed the potentially promising contribution of HTPS to the attainment of SDG-2 (see also Lindhout et al., 2017). Moreover, they emphasised that their core business (i.e. ...
Full-text available
In diploid potato the production of unreduced gametes with a diploid (2n) rather than a haploid (n) number of chromosomes has been widely reported. Besides their evolutionary important role in sexual polyploidisation, unreduced gametes also have a practical value for potato breeding as a bridge between diploid and tetraploid germplasm. Although early articles argued for a monogenic recessive inheritance, the genetic basis of unreduced pollen production in potato has remained elusive. Here, three diploid full-sib populations were genotyped with an amplicon sequencing approach and phenotyped for unreduced pollen production across two growing seasons. We identified two minor-effect and three major-effect QTLs regulating this trait. The two QTLs with the largest effect displayed a recessive inheritance and an additive interaction. Both QTLs co-localised with genes encoding for putative AtJAS homologs, a key regulator of meiosis II spindle orientation in Arabidopsis thaliana . The function of these candidate genes is consistent with the cytological phenotype of mis-oriented metaphase II plates observed in the parental clones. The alleles associated with elevated levels of unreduced pollen showed deleterious mutation events: an exonic transposon insert causing a premature stop, and an amino acid change within a highly conserved domain. Taken together, our findings shed light on the natural variation underlying unreduced pollen production in potato and will facilitate interploidy breeding by enabling marker assisted selection for this trait.
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The third most important food crop worldwide, potato (Solanum tuberosum L.) is a tetraploid outcrossing species propagated from tubers. Breeders have long been challenged by polyploidy, heterozygosity, and asexual reproduction. It has been assumed that tetraploidy is essential for high yield, that the creation of inbred potato is not feasible, and that propagation by seed tubers is ideal. In this paper, we question those assumptions and propose to convert potato into a diploid inbred line-based crop propagated by true seed. Although a conversion of this magnitude is unprecedented, the possible genetic gains from a breeding system based on inbred lines and the seed production benefits from a sexual propagation system are too large to ignore. We call on leaders of public and private organizations to come together to explore the feasibility of this radical and exciting new strategy in potato breeding. © Crop Science Society of America | 5585 Guilford Rd., Madison, WI 53711 USA All rights reserved.
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Key message: This is the first report of the production and use of a diploid inbred line-based F2 population for genetic mapping in potato. Potato (Solanum tuberosum L.) is an important global food crop, for which tetrasomic inheritance and self-incompatibility have limited both genetic discovery and breeding gains. We report here on the creation of the first diploid inbred line-derived F2 population in potato, and demonstrate its utility for genetic mapping. To create the population, the doubled monoploid potato DM1-3 was crossed as a female to M6, an S7 inbred line derived from the wild relative S. chacoense, and a single F1 plant was then self-pollinated. A genetic linkage map with 2264 single nucleotide polymorphisms was constructed and used to improve the physical anchoring of superscaffolds in the potato reference genome, which is based on DM1-3. Segregation was observed for skin and flesh color, skin and flesh pigment intensity, tuber shape, anther development, jelly end, and the presence of eye tubers instead of normal sprouts. Using the R/qtl software, we detected 10 genes, 7 of which have been previously mapped and 3 for which this is the first publication. The latter category includes tightly linked genes for the jelly end and eye tuber traits on chromosome 5. The development of recombinant inbred lines from this F2 population by single-seed descent is underway and should facilitate even better resolution of these and other loci.
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From 2006 through 2015, a research project on Durable Resistance in potato against Phytophthora (DuRPh) was carried out at Wageningen University and Research Centre. Its objective was to develop a proof of principle for durable resistance against late blight by cisgenesis. This public-funded project aimed at stimulating research on genetic modification and public debate on innovative genetic techniques. It was decided to clone and transfer late blight resistance (R) genes of crossable wild potato species (cisgenes) by Agrobacterium tumefaciens-mediated transformation without non-potato genes. A stack of multiple R genes were planned to be inserted into established varieties, thereby creating a dynamic variety in which the composition of the stacks may vary over space and time. Cisgenic plants were selected based on the expression of all inserted R genes and trueness-to-type. Within the project, 13 R genes from wild potato species were genetically mapped and three of them were cloned. Four varieties were transformed with one to three R genes. This was initially done using kanamycin resistance provided by a selectable marker gene of synthetic origin in order to quickly test the performance and stability of the introduced R genes and stacked R gene combinations. Once the functioning thereof was confirmed, marker-free transformations were conducted; thus, true cisgenic events were selected. The results about the different R genes, their chromosomal location, their specificity, the background dependence, the maximum size of a stack, its regeneration time and associated somaclonal variation frequency and its stability were studied. After selection and characterisation in the laboratory, the best cisgenic events were assessed in field trials for late blight resistance. This showed that inserted R genes were capable of turning a susceptible variety into a resistant one. Maximising longevity of the resistance was assured through resistance management research. It was shown that stacking of multiple R genes and monitoring how to deploy these stacks spatially and temporally could reduce fungicide use by over 80%. Communications through media and field demonstrations were manifold to allow public and policymakers to decide if cisgenesis is an acceptable tool to make potato farming more sustainable. Future deployment of the DuRPh strategy will depend largely on its status as a genetically modified crop or its exemption thereof. Worldwide near eradication of late blight would increase global annual potato production by close to 80 million tons, thereby contributing considerably to the needed additional global future food supply.
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Capsinoids have similar biological effects as capsaicinoids, including anticancer and anti-obesity properties. The Capsicum chinense ‘SNU11-001’ variety of pepper was previously reported to contain high levels of capsinoids due to a mutation in the putative aminotransferase (pAMT) gene, which leads to production of the capsinoid precursor vanillyl alcohol. Here, to develop Capsicum annuum pepper varieties with high capsinoid contents, marker-assisted backcrossing was performed during backcross breeding. For foreground selection, plants carrying the pAMT/pamt genotype were selected from BC1F1 and BC2F1 populations using SCAR markers derived from the unique pamt mutation of ‘SNU11-001.’ To obtain background selection markers, 412 single nucleotide polymorphism (SNP) markers were screened to identify SNP markers polymorphic between the ‘Shinhong’ paternal lines and ‘SNU11-001.’ Of the 412 SNP markers, 204 polymorphic SNP markers evenly distributed in pepper genome were selected. BC1F1 and BC2F1 plants carrying the pAMT/pamt genotype were subjected to background selection using these SNP markers. Multiple genotypes were analyzed using a high-throughput genotyping system. As a result, one BC1F1 plant 84 % similar to the recurrent parent and seven BC2F1 plants showing more than 96 % recovery of the recurrent parent genotype were selected. Genetic backgrounds of the selected BC2F1 plants were evaluated using the genotype-by-sequencing (GBS) method to confirm the background selection results from the SNP marker set. GBS results showed that the recovery rate and positions of introgressed segments were well matched between two methods, demonstrating MABC can be successfully performed with only ~200 SNP markers.
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This paper describes the progress that has been made since the draft genome sequence of potato has been obtained and the analyses that need to be done to make further progress. Although sequencing has become less expensive and read lengths have increased, making optimal use of the information obtained is still difficult, certainly in the tetraploid potato crop. Major challenges in potato genomics are standardized genome assembly and haplotype analysis. Sequencing methods need to be improved further to achieve precision breeding. With the current new generation sequencing technology, the focus in potato breeding will shift from phenotype improvement to genotype improvement. In this respect, it is essential to realize that different alleles of the same gene can lead to different phenotypes depending on the genetic background and that there is significant epistatic interaction between different alleles. Genome-wide association studies will gain statistical power when binary single nucleotide polymorphism (SNP) data can be replaced with multi-allelic haplotype data. Binary SNP can be distributed across the many different alleles per locus or may be haplotype-specific, and potentially tag specific alleles which clearly differ in their contribution to a certain trait value. Assembling reads from the same linkage phase proved to allow constructing sufficiently long haplotype tracts to ensure their uniqueness. Combining large phenotyping data sets with modern approaches to sequencing and haplotype analysis and proper software will allow the efficiency of potato breeding to increase.
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Key message: A 20K SNP array was developed and a comprehensive set of tetraploid cultivar was genotyped. This allowed us to identify footprints of the breeding history in contemporary breeding material such as identification of introgression segments, selection and founder signatures. A non-redundant subset of 15,138 previously identified SNPs and 4454 SNPs originating from the SolCAP project were combined into a 20k Infinium SNP array for genotyping a total of 569 potato genotypes. In this study we describe how this SNP array (encoded SolSTW array) was designed and analysed with fitTetra, software designed for autotetraploids. Genotypes from different countries and market segments, complemented with historic cultivars and important progenitors, were genotyped. This comprehensive set of genotypes combined with the deliberate inclusion of a large proportion of SNPs with a low minor allele frequency allowed us to distinguish genetic variation contributed by introgression breeding. This "new" (post 1945) genetic variation is located on specific chromosomal regions and enables the identification of SNP markers linked to R-genes. In addition, when the genetic composition of modern cultivars was compared with cultivars released before 1945, it appears that 96 % of the genetic variants present in those ancestral cultivars remains polymorphic in modern cultivars. Hence, genetic erosion is almost absent in potato. Finally, we studied population genetic processes shaping the genetic composition of the modern European potato including drift, selection and founder effects. This resulted in the identification of major founders contributing to contemporary germplasm.
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Vegetable production has been a major and dynamic activity devised by diverse human cultures to sustain their livelihood for centuries. Vegetables, being several times more productive per unit area than cereals, can play a vital role in facing food security and nutrition challenges in the coming decades. However, the predicted climate change and increased demand on limited land and water resources makes water conservation a key component of vegetable production systems. At the same time, there is an increased global demand for healthy and nutritious vegetables. Dramatic improvements have been achieved through breeding for important abiotic stresses and quality traits in many vegetables. Thus, successful emerging small or large commercial farmers now apply integrated strategies from farm to table, including planting, grafting, irrigation, use of modern cultivars and innovative marketing tools. In this chapter we highlight some technological advances in vegetable production, with emphasis on stand establishment and irrigation management for water-limited areas. We discuss the impact of breeding and genetics on the improvement of abiotic stress tolerance and provide evidences on the use of improved germplasm and cultivars to enhance the quality of vegetables. Finally, we discuss the critical role of marketing and consumer trends for vegetable products. © 2014 Springer Science+Business Media Dordrecht. All rights reserved.
India is the second largest producer of potato in the world. The Indo-Gangetic plains (IGP) is the main potato growing region accounting for almost 85% of the 1.8 Mha under the crop in India where it is grown as an irrigated crop during the winter season. Since IGP is in sub-tropical plains, duration of the thermally suitable window is the main determinant limiting yields. Hence the impact of climate change on potato in the IGP was assessed using MIROC HI.3.2 A1b and B1, PRECIS A1b, A2, B2 scenarios and estimated the potential adaptation gains. The potato crop duration in the IGP is projected to decrease due to climate change. The evapotranspiration (ET) is projected to increase while the water use efficiency (WUE) for potato yield is projected to decline in future climates as a consequence of low threshold temperatures for decline in WUE and yield than the ET. Results indicate that the upper threshold for ET decrease is ~23 °C while that for WUE is 15 °C. The optimal temperatures for tuber yield is ~17 °C and thus the reduction in WUE in future climates is discernable. Climate change is projected to reduce potato yields by ~2.5, ~6 and ~11% in the IGP region in 2020 (2010-2039), 2050 (2040-2069) and 2080 (2070-2099) time periods. Change in planting time is the single most important adaptation option which may lead to yield gains by ~6% in 2020 and its combination with improved variety or additional nitrogen may be required to adapt to climate change leading to positive gains by ~8% in 2020 and by ~5% even in 2050. However, in 2080 adoption of all the three adaptation strategies may be needed for positive gains. Intra-regional differences in the impact of climate change and adaptation gains are projected; positive impact in northwestern IGP, gains in Central IGP with adaptation and yield loss in eastern IGP even with adaptation.
In Kenya potato is an important food crop, second after maize. Potatoes are grown on 128,000 ha per year with average yields of about 8 tonnes per ha. The yield is far below its po-tential and should be improved to enhance food security. Of all potato growers in Kenya, 98% are characterised as small-scale farmers, producing less than 0.4 ha of potatoes per year per farm (total of two planting seasons). They produce 83% of the national production. Lack of good quality seed is a main reason for low yields. The fast-track seed project aims to improve availability of certified seed. Imported seed needs to be multiplied locally once or twice to reduce the cost price and make it affordable for small-scale growers. It is estimated that there is potential for at most 1,000 ha of professional seed production in Kenya to meet the market demand for certified seed. In Kenya about 98% of ware potatoes are sold for 'fresh' consumption. For industrial processing the most favourable opportunities are the production of crisps and fresh - not frozen - French fries. To change the low-input low-output strategy of many small-scale farmers linkages between producers and markets need to be improved. In the short term, improvement of potato growing and crop management should receive higher priority than storage and mechanisation. Storage and mechanisation should be adapted to local conditions and needs. Two cases have been indicated in which investments in professional cooled storage facilities are necessary; - storage of seed potatoes of varieties with a long dormancy period; - storage of ware potatoes for processing industry and some high-end retailers. Long-term seed storage in modern stores required for varieties with long dormancy increases the cost price of seed by 50% compared with short-term storage in diffused light stores. This makes short dor-mancy an important trait for potato varieties in Kenya, where potatoes are planted during two growing seasons. Machinery supplied for potato growing in Kenya should be geared to local needs. Higher yields and more multiplications of imported seed reduce the cost price of seed potatoes produced in Kenya. The cost price of imported seed after two multiplications is competitive with locally produced seed from minitubers after three multiplications. This report describes the current situation of the potato sector in Kenya and opportunities for further de-velopment. The study focuses on possibilities of the Dutch agribusiness to facilitate these developments. Existing information has been collected, although information on potato production was not easily availa-ble. In addition, during a mission to Kenya stakeholders were visited and interviewed. This resulted in cost- price calculations for seed potato production (short and long-term storage, imported seed and minitubers) and market information.