1. Introduction
Vegetable brassicas
are an important and highly diversified group of crops grown world-wide
that belong mainly to the species Brassica oleracea and B. campestris. In
the western hemisphere, including Europe, the predominance goes to B. oleracea,
a species that includes important crops such as cabbage, broccoli, cauliflower,
Brussels sprouts, etc. In Asia, B. campestris is the most cultivated species
owing to the great importance of Chinese cabbage. Turnip and turnip greens,
which also belong to B. campestris, are cultivated world-wide but have much
less economic importance. B. juncea, that includes the vegetables mustards,
has some economic relevance in far eastern countries but is a minor crop
at the world level. B. napus is probably the brassica receiving the most
attention from breeders at the world level, especially in Canada and France,
but it is only used for rapeseed oil. Therefore, it will not be considered
in this survey.
The centre of origin
of B. oleracea is the Mediterranean region. Vegetable brassicas are cultivated
in Europe since very ancient time from where they have spread to other parts
of the world (Nieuwhof, 1969). Brassicas include many different morphotypes,
are well adapted to temperate climates, require quite simple cultivation
techniques and produce abundant and nutritious food for man and domestic
animals. In the cold-winter regions of Europe white cabbage is processed
as sauerkraut, while in the mild-winter regions, brassicas are the most
important vegetable during the cool season. In Korea, Chinese cabbage is
used to prepare "Kim chi", the very popular national preserve.
The versatility of uses
and the adaptability to growing systems and man's life-style explain the
importance brassicas have had for centuries. However, the increasing consumption
of salad vegetables, the availability of a highly-diversified all year round
offer of imported and greenhouse grown vegetables make brassicas less attractive
for consumers in developed countries. Where world trade tends to dominate
the vegetable market, local availability is no longer a comparative advantage
for brassicas. However, vegetable brassicas have new appeal in developed
countries due to the potential for market diversification of some new crop
types, and to the use of cauliflower and broccoli as salad crops. With an
increasing number of consumers aware of the importance of diet on human
health, the potential benefits of a diet with green vegetables have given
brassica vegetables a better image during recent years. In less developed
countries brassicas are still considered as an important source of cheap
and abundant food.
Breeding strategy and
targets are dependent on market trends. Successful breeders anticipate changes
in the market by developing new varieties that are ready to be released
to the growers when their demand increases. It is therefore interesting
to see how breeding is reacting to eventual changes in brassica consumption
and to evaluate the potential influence that the brassica market and growing
systems may have on the definition of breeding targets and priorities.
The objective of this
paper is to analyse the present situation and the perspectives of brassica
breeding based on the authors personal knowledge of the sector and on information
collected through an international survey of brassica breeders. The breeding
of F1 hybrid winter-cauliflower in Brittany is presented as an example of
successful co-operative breeding in tune with both the demands of growers
and the markets alike.
2. Survey on brassica breeding
2.1 Questionnaire
A questionnaire was
mailed to 26 private and public brassica breeders world-wide, in January,
1998. The questions included the number of breeders per company/institution,
the changes in the number of breeders during the last ten years, the perspectives
for brassica breeding and its relative importance to other vegetable crops,
the relative distribution of breeding activity per crop, the ranking of
breeding objectives and the three major advantages/strengths and threats/weaknesses
for brassica breeding in the near future.
A total of 18 replies
were received from 16 private and 2 public breeders. Thirteen replies were
from Europe, two from Japan and one each from USA, India, and Korea.
2.2 Breeding activity
There is a total of
70 brassica breeders, outside Korea, working for the companies/institutions
questioned. The number of breeders per company/institution varies from 1
to 12. In 60% of cases there were less than five breeders per company/institution.
Korea is an exception with approximately 60 brassica breeders, from which
40 work in seed companies, 10 in research stations and 10 in universities.
This breeding activity reflects the enormous importance of brassica crops
in this country.
The number of breeders
increased during the last ten years in 11 out of the 18 companies/institutions
and has not changed in 7. Although, when the same group of persons were
asked about the future, 9 said that brassica breeding will increase, 8 replied
that it will not change and one said that it will decrease. This is a situation
of moderate optimism. Brassica breeding is not likely to decrease, rather
it will remain stable or increase slightly.
Korea is again the exception.
Breeding has been stable during the past ten years and is likely to decrease
in the future. This is explained by the tendency for a decrease in the consumption
-currently extremely high- of brassicas due to dietary diversification,
and also to substitution of locally-bred varieties by imported F1 hybrids.
2.3 Relative importance of brassica crops 
The companies/institutions
showed big differences in the pattern of distribution of breeding activity
between crops. The crops deserving the most breeding attention vary considerably
between company/institution . There is a tendency for specialisation of
companies in a reduced number of crops. In 10 companies/institutions over
70% of the breeding activity is concentrated in one or two crops. In six
companies/institutions one single crop may absorb from 70 to 100% of the
breeding activity. Some small breeders become very specialised e.g. three
breeders dedicate 80% of their activity to cauliflower and one breeder 100%
to kohlrabi. There are still several less specialised companies/institutions
where the distribution of breeding activity is more equitable and tends
to follow the relative economic importance of the various crops. In Korea
60% of the brassica breeding is on Chinese cabbage.
Cauliflower is the most
important crop attracting over 50% of the breeding activity in 7 breeding
companies/institutions (Table 1). Only two companies/institutions do not
have cauliflower breeding. Broccoli and white cabbage follow in decreasing
order of importance. Savoy cabbage and Brussels sprouts are much less important.
Table 1. Number of companies/institutions per class of percentage
of breeding activity dedicated to a certain crop
| |
Number of responders per class of importance
|
| |
High
|
Medium
|
Low
|
| Uniformity |
15
|
2
|
1
|
| Disease resistance |
13
|
5
|
0
|
| Appearance |
12
|
4
|
2
|
| Crop yield |
10
|
|
1
|
| Pest resistance |
3
|
9
|
6
|
| Nutritional quality |
1
|
9
|
8
|
| New crop types |
1
|
4
|
13
|
2.4 Breeding objectives 
Breeding objectives
can be addressed to satisfy the grower or the consumer and be considered
in terms of crop improvement and product improvement. Such a division of
objectives, that may seem artificial since growers cannot survive by using
varieties that are not accepted by the market, can be used with advantage
for discussing breeding strategy and evolution. The main criteria for crop
improvement are yield, disease resistance or abiotic stress, uniformity
and continuity of cropping. Breeding for appearance, commercial quality,
shelf life, taste, and nutritional value is part of product improvement.
The most important breeders
objective in our survey is crop uniformity which received 15 citations as
high priority (Table 2). A uniform brassica field makes grading much easier
and reduces harvest time. The final objective is to have a single-harvested
field of uniform quality. High uniformity has been almost impossible to
achieve with open-pollinated varieties owing to the cross-pollination habit
of brassicas. The introduction of F1 hybrids, that can produce a genetically-uniform
population, has progressed slowly. Until recently brassica hybrid breeding
has been using the sporophytic self-incompatibility mechanism since there
is no cytoplasmic male sterility in B. oleracea. The instability and complex
inheritance of the self-incompatibility mechanism makes its use difficult
and conducive to low quality F1 hybrids.
However, the production
of brassica F1 hybrids is now developing faster, albeit with some technical
difficulties, using double-haploid parent lines obtained through microspore
culture and cytoplasmic male sterility introduced from Raphanus sativus
into B. oleracea.
Table 2. Number of companies/institutions per class of importance
for each breeding objective
| |
Number of responders per class of importance
|
| |
High
|
Medium
|
Low
|
| Uniformity |
15
|
2
|
1
|
| Disease resistance |
13
|
5
|
0
|
| Appearance |
12
|
4
|
2
|
| Crop yield |
10
|
7
|
1
|
| Pest resistance |
3
|
9
|
6
|
| Nutritional quality |
1
|
9
|
8
|
| New crop types |
1
|
4
|
13
|
Disease resistance is
also a very important breeding objective ranked as first priority by 13
out of 18 responders and not considered a low priority by anyone. Pest resistance
is much less important since most responders considered it a medium or low
priority (Table 2) 
Sources of resistance
to important diseases e.g. clubroot (Plasmodiophora brassicae), black rot
(Xanthomonas campestris pv. campestris), fusarium yellows (Fusarium oxysporum
f. conglutinans) and downy mildew (Peronospora parasitica) were identified
(Chiang et al., 1993) but have not been widely transferred into commercial
varieties yet. The relative delay of breeding for pest and disease resistance
may be explained by the low destructive effect of pathogens in brassicas
and the need to incorporate the resistant genes into a high number of crop
types. There is no disease that is a serious limitation to brassica growing
in large areas and the most destructive diseases, for example clubroot,
can either be controlled with chemicals or kept below the threshold of economic
damage with relevant growing practices. The economic return of investment
in breeding for disease and pest resistance is quite low because it is dispersed
among too many different crop types, and resistant varieties are competing
directly with non-resistant ones that can still be used by the growers without
too much problem. The high interest in and the increasing present demand
for breeding for pest and disease resistance is related to a generalised
interest in releasing "environmentally friendly" varieties requiring
a sparse use of chemicals.
Appearance, including
colour and shape, is another important trait (Table 2) and the only major
breeding objective that is addressed to the consumers. The rapid transformation
of the vegetable market with the improvement of packing and display facilities,
the large offer of commodities all the year round, and the increased presence
of colourful and appealing fruit and salad vegetables has forced brassica
producers to rise their presentation and quality standards. Growers are
looking for high commercial quality, including adequate size and shape,
good colour, firmness, and appearance to have the produces easily accepted
by the trade.
Crop yield, the traditional
first priority for breeders, ranks only fourth among breeding priority (Table
2), confirming the high relative importance of qualitative traits in vegetable
breeding.
Nutritional quality
has as yet little importance since it was ranked as medium or low priority
by 17 out of 18 responders (Table 2). A similar low interest was shown by
breeding for new crop types. It is surprising to see that crop diversification
and nutritional quality, both of which are considered to be important marketing
advantages for the future, have received so little attention from brassica
breeders. Perhaps this is because crop diversification has long been a breeding
objective in brassicas (as well as in other vegetable crops) without being
accepted as such. The best known example is broccoli which was introduced
from Italy to the rest of Europe and the United States and has now become
one the most important brassica crops world-wide. Chinese cabbage, due to
its increasing cultivation and consumption in western countries, is also
a good example of crop diversification. Lesser known brassicas e.g. romanesco
cauliflower, mustard greens or kohlrabi that could also be interesting sources
of crop diversification do not, as yet, receive the same attention.
2.5 Advantages and weaknesses of brassica breeding 
The responders were
asked to cite the three most important strengths and weaknesses of brassica
breeding in the near future. Those items mentioned by more than one responder
are listed in Table 3. The answers were quite diverse, with little agreement
about the likely perspectives for vegetable brassica breeding. What is considered
an advantage for some breeders may be a threat for others. This undoubtedly
arises from the wide diversity of crops and commercial situations concerned.
The potential advantages
of the application of biotechnology to brassica breeding received the largest
agreement among responders (Table 3). This is also true for vegetable breeding
in general, but in the case of vegetable brassicas is more prominent due
to the direct impact received from the huge research on Arabidopsis thaliana
and B. napus. Breeders working with B. oleracea can have easy access to
the knowledge on the use of molecular markers, identification of genes for
resistance, transformation, etc. developed for A. thaliana and B. napus.
There is no agreement
on the impact of the market on brassica breeding. Seven responders considered
the market value and consumer acceptance of brassicas as an advantage while
8 responders anticipate adverse market perspectives owing to changes in
consumer preferences. The good market perspectives are related to the potential
benefits of brassicas for human health and the increased awareness of consumers
about the role of the diet on disease prevention. It may be expected that
in some more advanced markets the consumption of broccoli and green vegetables
will increase thanks to their role in protecting against human cancer. Also,
the good adaptability of brassicas to a wide range of growing conditions
could encourage their use as a cheap source of abundant and nutritious food
in developing countries where brassicas are not traditional. However, it
is difficult to say if these market advantages can balance the overall tendency
for the decrease in consumption in developed countries where brassicas tend
to be replaced by the more appealing fruit and salad vegetables.
The slow progress and
difficulties of hybrid breeding pointed out by five responders are related
to the use of sporophytic incompatibility for the production of F1 hybrids.
However, the progress in the use of cytoplasmic male sterility will make
hybrid breeding much easier in the near future.
A last comment on the
concern of five responders about the impact that the fusion policy and reduced
profitability in the seed industry may have on the number of brassica breeders.
It is evident that the concentration of seed companies will merge or cancel
some breeding programmes to reduce costs. Then there will be fewer brassica
breeders and the growers will be dependent on a narrower genetic background.
Strengths and weaknesses
of brassica breeding mentioned by more than one responder *
| Strengths |
Weaknesses |
Wide application of biotechnology to brassica breeding
e.g. double-haploidization, genetic transformation (11)
Market value and consumer acceptance (7)
Increasing awareness of the high importance of brassicas in human
diet (7)
Potential use of genetic diversity (5)
Disease and pest resistance (3)
Adaptability to a wide range of growing conditions (3)
Crop yield (2)
|
Slow progress on breeding for disease and pest resistance
e.g. blackrot, clubroot (9)
Adverse market perspectives owing to changes in consumer preferences
(9)
Fusion policy and reduced profitability in the seed industry (5)
Slow progress and difficulties of hybrid breeding (5)
Nutritional value and taste (4)
|
* The number of responders per item is presented in brackets
3. Winter-cauliflower breeding in Brittany 
Brittany is the leading
region for vegetable crop production in France with 25% of the national
output and one of the most important within Europe. Major regional crops
include cauliflower, artichoke, shallots and broccoli (80% of the national
production for these four). Field cropping of vegetables occupies some 70,000
ha, largely in the north coastal area where good soils and a favourable
maritime climate allow year-round production.
The growers are organised
into marketing co-operatives, themselves co-ordinated through a regional
body, the CERAFEL (Comité Economique Agricole Régionale Fruits
et Légumes de Bretagne). This organisation of the market, going back
some 30 years, coupled with the development of rapid transport links has
stimulated exportations to those countries of Europe where production of
vegetable crops in the winter months is impossible for climatic reasons.
Today, more than 50% of the area of brassica crops is exported.
Financial support by
the industry (via the CERAFEL) for scientific and technical work has focused
largely on the genetic improvement of cauliflower, the region major vegetable
crop. Strong links have been forged with the INRA (Institut National de
Recherche Agronomique) and a veritable network of research and development
organisations has grown up across the region. There are currently 30 research
and technical people directly involved in the vegetable brassica breeding
programmes at this moment.
One such organisation
is the OBS, l'Organisation Bretonne de Sélection. Established in
1970 through a joint initiative of the local grower co-operative, the OBS
has the particularity of being a wholly grower-owned private company dedicated
to crop improvement for a vegetable producing region (admittedly one of
Europe's largest). In this sense, the company is a means to an end and plant
breeding just one aspect of an overall objective: total control of crop
quality. For winter cauliflower, the main objective is to provide growers
with a complete range of locally-adapted F1 hybrids and , for this crop,
70% of the demand for seed within Brittany is currently met by the OBS.
This can be considered
something of a success story when one considers that thirty years ago all
of the cauliflower seed in the region was farm-saved. The need to improve
and homogenise the quality of the crop was the deciding factor in the creation
of the OBS and the ties subsequently developed with INRA.
The identification and
the multiplication of the best of the farm-saved stocks was the initial
task of the OBS. Varieties were maintained and improved using mass selection
with progeny testing, and with seed production under polyethylene tunnels.
These techniques allowed a greater number of growers access to the better
open pollinated varieties, so contributing to the qualitative and quantitative
progression of the crop in the area.
The major drawbacks
of this approach were an absence of control over the breeding system and
the difficulty of maintaining parental clones from one generation to the
next. With the emergence of reliable tissue culture techniques in the early
1980's, this latter obstacle was removed, so permitting a progression towards
more stable open-pollinated varieties of the "synthetic hybrid"
type.
In parallel to this
improvement of open-pollinated material, hybrid breeding programmes for
both autumn and winter cauliflower were started in 1970, jointly funded
by INRA and the CERAFEL, with scientific impute form the INRA focused on
the understanding and the exploitation of the two breeding systems available
in brassicas, self-incompatibility and male sterility.
The main breeding criteria
for cauliflower crop improvement (satisfaction for the grower) are yield,
uniformity and continuity of cropping. The main criterion for product improvement
(satisfaction of the consumer) is curd quality: whiteness, structure, freedom
from disease and nutritional and sensorial value.
At OBS, breeding efforts
are currently focused on providing F1 hybrids for the entire cropping calendar,
but increasing consumer awareness and concern for environmental issues has
put disease resistance/tolerance high on the scale of priorities.
Genotype x environment
effects are great in these crops, and importance is attached to the screening
of prototype hybrids. Any hybrid reaching the stage of large scale seed
production at the OBS will have been evaluated over at least 4 different
years and on up to 25 different sites across northern Brittany. The final
stages of evaluation include commercial-scale grower trials. All regionally-bred
material is compared to varieties available from other seed companies, both
on the regional experimental stations and on growers' holdings.
Given the considerable
breeding effort in this crop, hybrid varieties will inevitably replace the
remaining open-pollinated populations within the next five or so year.
Acknowledgements
The authors want to
thank the helpful co-operation of the responders to the questionnaire: Peter
Crisp, Crisp Innovar Ltd, UK; S. Kennedy, Elsoms Seeds Ltd, UK; A J M van
der Nieuwenhuizen, Ryk Zwaan, The Netherlands; Erling Hegelund, Dæhnfeldt,
Denmark; S. Jennings, A.L. Tozer Ltd, UK; C.M. Binnendijk, Enza Zaden B.V.,
The Netherlands; S.K. Tripathi, Mahyco Seeds, India; Remi Levieil, Limagrain,
France; P. Tjeertes, Novartis Seeds B.V., The Netherlands; Hiromasa Noguchi,
Sakata Research and Development Center, Japan; A. Blom, Huizer Zaden Holland
B.V., The Netherlands; Paul Degreef, Sakata Seed Europe B.V., The Netherlands;
M. Dickson, Cornell University, USA; Jung-Myung Lee, Kyung Hee University,
Korea; Koji Sakamoto, Takii Plant Breeding & Exp. Station, Japan; Nunhems
Zaden B.V., The Netherlands; Frans van der Bosch, Royal Sluis, The Netherlands;
T. Lunn, OBS, France.
References
Chiang, M.S, C. Chong,
B.S. Landry and R. Crête. Cabbage in G.Kalloo and B.O.Bergh (eds.).
Genetic Improvement of Vegetable Crops. Pergamon Press, Oxford, 1993
Nieuwhof, M. Cole crops.
Leonard Hill, London, 1969
|