International Journal of Vegetable Science

ISSN: 1931-5260 (Print) 1931-5279 (Online) Journal homepage: http://www.tandfonline.com/loi/wijv20

Breeding Cauliflower: A Review

B.K. Singh, Bijendra Singh & P.M. Singh

To cite this article: B.K. Singh, Bijendra Singh & P.M. Singh (2017): Breeding Cauliflower: A
Review, International Journal of Vegetable Science, DOI: 10.1080/19315260.2017.1354242

To link to this article: http://dx.doi.org/10.1080/19315260.2017.1354242

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Download by: [Cornell University Library] Date: 22 July 2017, At: 00:47
 Breeding Cauliflower: A Review

B.K. Singh, Bijendra Singh and P.M. Singh

Indian Council of Agricultural Research-Indian Institute of Vegetable Research, Shahanshahpur-

221305, Varanasi, Uttar Pradesh, India

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Abstract

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Cauliflower (Brassica oleracea L. var. botrytis L.) is continually being improved to

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increase sustainability of production, enhance nutritional quality and reduce waste. Development

of resistances against abiotic (high temperature and rainfall) and biotic stresses (diseases and

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insects) is necessary for this purpose. Breeding plays a vital role in addressing these issues

through development of superior varieties/hybrids. Globally, more than 20,000 accessions of C-

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genome taxa of B. oleracea are maintained in >100 gene-banks, European countries host the
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world's largest and most diverse collections. The favorable genes and quantitative trait loci

(QTL) for various traits of economic importance are scattered across cultivated types, breeding
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lines and wild populations (i.e., primary, secondary, tertiary, and other gene pools) of B.

oleracea, with efforts being made to integrate 2- or multi-tiered breeding approaches for
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broadening the genetic base and introgressing genes and QTL of resistance, quality and

productivity into elite backgrounds. The F1 hybrids that largely replace open-pollinated varieties,
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offer desirability, uniformity and consistency of vigor, yield and quality; but their commercial
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seed production face technical difficulties which can be overcome by developing a robust system

of self-incompatibility (SI), Ogura cytoplasmic male sterility (CMS) and doubled haploid (DH)

parental lines, as well as proper management of pollinators. The rapidly growing knowledge of

advance tools of parental line development, molecular markers and biotechnological techniques

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will provide enhanced precision and extend options to support future cauliflower breeding

programs.

Keywords: Brassica oleracea; cytoplasmic male sterility; molecular marker; self-

incompatibility

Cauliflower (Brassica oleracea L. var. botrytis L.) has wide adaptability from temperate

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regions to the tropics. It has 18 diploid and somatic chromosomes (2n = 2x = 18). The word cole

is variously spelled as Kale (English), Kohl (German), Kool (Dutch), Kal (Scandinavian), Kaali

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(Finnish), Kaol and Kol (Breton), Chou (French), Col (Spanish), Cal (Irish), Cavolo (Italian),

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and Couve (Portuguese); but the word cole is more recognized in the literature worldwide

(Chatterjee and Kabir, 2002). The cole crops are a group of highly differentiated Brassica plants
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grown all over the world. Globally, cauliflower grows best between the latitudes 11-60°N with

average temperature ranging from 5-8°C to 25-28°C. Cauliflower may tolerate temperature from
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-10°C to 40°C for a few days during the vegetative growth period.

Area, production and productivity of cauliflower in the world, and India, is 1258 and
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433.9 thousand ha, 22840 and 8573 thousand Mt, and 18.2 and 19.8 Mt·ha-1, respectively
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(Anonymous, 2014). The top 10 producing countries are China (35.77%), India (34.49%),

Mexico (2.13%), France (1.56%), Italy (1.36%), Poland (1.23%), USA (1.16%), Pakistan
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(1.07%), Germany (0.53%) and Egypt (0.48%). The important Indian states producing

cauliflower are West Bengal, Bihar, Maharashtra, Madhya Pradesh, Odisha, Gujarat, Haryana,
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Chhattisgarh, Jharkhand, Assam and Uttar Pradesh (Anonymous, 2014). The secondary

metabolites of Brassica are indole-3-carbinol (I3C), indole-3-acetonitrile (IAN), 3-3-

diindolylmethene (DIM), 1-methoxyindol and 3-carbaldehyde which can induce cytochrome P-

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450 dependent mono-oxygenase activity and protect against cancer and radiation in mice

(Bradfield and Bjeldanes, 1987; Fang et al., 2013).

Origin, evolution and domestication

Detailed studies on molecular phylogenetics, diversification analysis, and historical

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biogeography (Arias et al., 2014) established that the tribe Brassiceae originated in the

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intersection between the Arabian Peninsula and North Africa (Saharo-Sindian region), about 24

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million years ago (Mya). The nigra lineage diverged from the oleracea/rapa lineage about 20

Mya with the former continuing to differentiate in southwestern parts of the Mediterranean

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region; while the latter diversified in the Saharo-Sindian region and colonized the European
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Mediterranean through the east. The ‘core oleracea’ clade, containing B. rapa and the B.

oleracea originated in the northeastern Mediterranean ca. 6.5 Mya, possibly at the time of the
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Messinian Salinity Crisis. At this time, the Mediterranean went through cycles of partial

desiccation lasting several hundred thousand years, reducing its extension, with salt deposits
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accumulating and new drier and open environments being created. According to Arias et al.

(2014), B. oleracea and its C-genome wild relatives diversified in the northeast Mediterranean
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region about 1.5 Mya and then spread through the rest of Europe. Wild primitive forms of B.

oleracea grow on cliffs of the Atlantic coast of Europe. Eventually, B. oleracea was brought to
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the east Mediterranean region where it became fully domesticated and gave rise to a wide range
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of cultivated forms. Taxonomic studies suggest that the progenitor of B. oleracea exists in 9

chromosomes in wild B. oleracea kale (Snogerup, 1980). Restriction Fragment Length

Polymorphism (RFLP) studies indicate that a primitive cultivated B. oleracea might have

evolved from wild B. oleracea (Song et al., 1988). Variation within, and between, subspecies of

B. oleracea, and present day cultivated cole crops evolved (Table 1) after a long time of

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natural/artificial hybridization, mutation, selection and domestication. All forms are descended

from a common kale like ancestor, the wild cabbage (B. oleracea L. var. sylvestris L.) which is

still prevalent in western and southern Europe, and North Africa.

Schulz (1919) anticipated that many cole crops are derived from the Mediterranean wild

forms of oleracea and suggested B. cretica as the probable progenitor of cauliflower. Boswell

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(1949) stated cauliflower originated in the Islands of Cyprus from the wild cabbage; moved to

Syria, Turkey, Egypt, Italy, Spain and north-western Europe; and is thought to have been

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domesticated in the Mediterranean region (Home, 1954; Helm, 1963; Nieuwhof, 1969). Hyams

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(1971) considered cauliflower was first noticed, selected and propagated in Syria. Recent

molecular and genetic studies provide evidence suggesting that cauliflower arose in southern
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Italy from a heading Calabrese broccoli via an intermediate Sicilian crop type (Smith and King,

2000). An herbalist, Dodens in 1578 presented the first description and illustration of cauliflower
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in cultivation in France around the 16th century, and available in the markets in England as early

as 1690. In the USA, cauliflower was first mentioned in 1806 and attained commercial status
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after 1920. In England, the cauliflower industry started in the early 19th century with Cornish

types, commonly grown in western Cornwall (Home, 1954; Johnstone, 1963). Cornish
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cauliflowers were grown until around 1920 when English housewives began preferring the
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white-curded ‘Roscoff’ from France. The ‘Roscoff’ was closely, and jealously, guarded, and

seed were not available. However, in 1924 its seed reached England and were multiplied for
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breeding (Johnstone, 1963). Later, ‘Cornish’ types were replaced by ‘Roscoff’ and other

improved strains (Swarup and Chatterjee, 1972).

According to the Germplasm Resources Information Network (GRIN) nomenclature,

there are several important cultivated B. oleracea groups (Table 2; Anonymous, 2015; Maggioni,

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2015). The cytogenetic relationships of the main species of genus Brassica have been depicted in

the U’s triangle (Figure 1) in which B. nigra (L.) Koch (n = 8, black mustard, genome-B), B.

oleracea L. (n = 9, cole crops, genome-C), and B. rapa L. (B. campestris in the past, n = 10,

turnip, genome-A) represent the 3 diploid species in the vertices; and altogether they developed 3

amphidiploid species by intercrossing, i.e., B. carinata A. Braun. (n = 17, Abyssinia mustard,

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genome-BC), B. juncea (L.) Czern. (n = 18, brown mustard, genome-AB), and B. napus L. (n =

19, oilseed rape, genome-AC). The creation of new species during the 1920s by Karpechenko

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who developed the synthetic genus Raphanobrassica by crossing of Raphanus sativus L. with B.

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oleracea L. var. capitata L. to combine their desirable traits (Karpechenko, 1924); and hence U’s

triangle has been extended by adding diploid R. sativus (n = 9, radish, genome-R) and
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amphidiplod Raphanobrassica (n = 18, raphanobrassica, genome-RC) (Figure 1). Since all

cultivated and wild taxa belonging to the same C-genome are grouped under the species B.
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oleracea, the cultivated B. oleracea are referred to as cole crops. They share the same

chromosome number (2n = 18) with a large number of wild species of Brassica usually found
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growing in the Mediterranean region, and on rocky limestone cliffs along the Atlantic coasts of

Britain, France and Spain (Song et al., 1990; Maggioni et al., 2010; Maggioni, 2015).
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Taxonomically, cole crops belong to the order Brassicales (Cruciales), family Brassicaceae
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(Cruciferae), tribe Brassiceae, subtribe Brassicinae, genus Brassica, section Brassica and species

oleracea. Pachytene chromosome studies have shown that the B. oleracea is a triple tetrasomic
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for chromosome types B, C and E comprising the genome formula ABBCCDEEF with 6 basic

genomes and showing some secondary pairing.

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Major types of cauliflower

European cauliflower: Systematic and extensive cultivation of cauliflower occurred first in Italy

where the ‘Originals’ or ‘Italians’ were developed. These ‘Original’ or ‘Italian’ types were taken

to France, England, Germany and the Netherlands where some important local types were

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developed such as the ‘Northerns’ in Yorkshire and Derbyshire; the ‘Cornish’ in Cornwall; the

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‘Angers’ and ‘Roscoff’ in Brittany; and the ‘Erfurt’ or its allied ‘Snowball’ in Germany and the

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Netherlands. Broadly, on the basis of suitability of growing conditions, the European

cauliflowers have been grouped into winter type, i.e., winter cultivation (‘Originals’ or ‘Italians’,

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‘Cornish’, ‘Northerns’, ‘Roscoff’, ‘Angers’) and summer type, i.e., summer cultivation (‘Erfurt’
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and ‘Snowball’). The chief characteristics of important cauliflower types are broadly elaborated

by Swarup and Chatterjee (1972).

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Indian cauliflower: The Indian/tropical cauliflowers grown in India are characteristically

different from European types, and are tolerant to high temperature and humid conditions, the
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earliest maturing types known, and do not require vernalization for bolting. Swarup and

Chatterjee (1972) hypothesized that Indian cauliflowers possessed most of the genes contributed
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by the ‘Cornish’ types and a few genes by ‘Roscoff’, ‘Italian’ and ‘Northern’ types. Certain

favorable genes for tolerance to high temperature and rainfall, unknown in European types, have
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been successfully utilized in developing the improved varieties Early Patna, Early Benaras and
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Early Market in India; Pua Kea in Hawaii; Campinus in Brazil; Improved Japanese and D 96 in

Israel; and Extra Early in Taiwan. In India, most cauliflower varieties are called by the name of

the month in which the curds become ready to harvest, i.e., Kunwari, Katki, Agahani, Pusi and

Maghi. These are highly heterozygous for maturity, plant growth and curd traits. The cauliflower

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varieties grown in India are classified into 4 groups, and the varieties grouped on the basis of

period of curd maturity (Table 3; Chatterjee and Swarup, 1983; Singh and Devi, 2015).

India experiences a wide range of weather conditions throughout its geographical regions.

In North Indian plains, the temperature differs drastically from Punjab to Bihar and affects crop

productivity if sowing/transplanting is not at the correct time. From 2013-2015, the range mean

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monthly temperature at the research farm of Indian Council of Agricultural Research-Indian

Institute of Vegetable Research (ICAR-IIVR) Varanasi, Uttar Pradesh, India was 26.3-34.4,

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22.0-31.8, 15.9-28.8 and 11.0-22.5°C, respectively throughout September, October, November

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and late-November to mid-December which is higher than have been experienced for the

respective periods, i.e., 20-27, 20-25, 16-20 and 12-16°C (Table 3). Realizing the critical role of
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temperature for curding, the varieties must be categorized on the basis of temperature

requirement rather than the period/month of maturity. Hence, varieties of Indian cauliflower
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should be reclassified as Extra Early (>30°C), Early (25-30°C), Mid (20-25°C), Mid-late (15-

20°C) and Late or ‘Snowball’ (<15°C) to avoid ambiguity, which appears to be on sound footing
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that can be easily understood and followed.

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History in India

Cauliflower was introduced to India by Dr. Jemson; a botanist at Kew Garden, London,
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and in-charge of Company Bagh (renamed as Government Botanical Garden) at Saharanpur,

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Uttar Pradesh in 1822. The Royal Agri-Horticultural Society, Calcutta, West Bengal introduced

seed of European vegetables from South Africa in 1824. Seed imported from England and South

Africa was given to growers in North India. From 1822-1929, these growers through selection,

unconsciously evolved present day Indian cauliflowers which have the ability to grow under high

temperature and humidity conditions, tropical in nature, with potential to produce seed in the

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north Indian plains. The first 4 Indian varieties listed by Sutton and Sons, India, in 1929 were

Early Crop Patna, Main Crop Patna, Early Crop Benaras and Main Crop Benaras. It is likely that

the Cornish type contributed most of the genes including long stalk, open growth habit (plant

type-2), and yellowish, uneven and strong flavored curds. Some leaves and curd characteristics

were contributed by Roscoff, Italian and Northern types. The present day Indian cauliflowers are

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recognized as different types (Nieuwhof, 1969; Swarup and Chatterjee, 1972; Crisp, 1982).

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Genetic resources

Genetic resources provide diversity necessary for genetic improvement. For B. oleracea,

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the primary genepool, including all taxa that easily produce fertile hybrids, can be represented by
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the C-genome taxa of the B. oleracea group. The secondary genepool, represented by the taxa

potentially capable of experimental hybridization with B. oleracea, is extended to the so-called

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Brassica coenospecies. The genetic resources are conserved as ex-situ (gene-banks) and as in-situ

populations (centre of origin, diversity and domestication or place of cultivation). Molecular

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resources are also increasingly useful, including expressed sequence tags (ESTs), full length

cDNA clones of many genes, and bacterial artificial chromosome libraries (Maggioni, 2015).
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With adoption of improved cultivars/hybrids, genetic variability in most cole crops, including

cauliflower, is disappearing from the public domain causing serious problem of genetic erosion.
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Worldwide, more than 20,000 accessions of C-genome taxa of B. oleracea are conserved in >100
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gene-banks of various countries (Table 4). In India, Indian Council of Agricultural Research-

Indian Agricultural Research Institute (ICAR-IARI) Regional Station, Katrain, Himachal

Pradesh; ICAR-IARI, New Delhi; ICAR-National Bureau of Plant Genetic Resources, New

Delhi; ICAR-Indian Institute of Vegetable Research, Varanasi, Uttar Pradesh; Dr. Yashwant

Singh Parmar University of Horticulture and Forestry, Solan, Himachal Pradesh; and some

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private seed companies are doing genetic improvement and maintaining germplasm of B.

oleracea. In Europe and Mediterranean regions, i.e., the center of origin, diversity and

domestication, the local forms of B. oleracea are still extensively grown in home gardens

indicating that the rich diversity is still present on-farm; but at the same time a comprehensive

inventory of on-farm diversity is not available. India too, home for evolution of tropical

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cauliflower, has a lot of genetic variability, yet its activities involving germplasm maintenance

on the international arena is minimal. It is critical that these resources are properly conserved,

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documented, and made accessible for use in breeding.

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Curd and its quality an
Botanically and morphologically, cauliflower curds are the ‘pre-floral fleshy apical

meristem’ which consists of a shoot system with short internodes, branch apices and bracts.
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Unlike other cole crops, most forms of Indian cauliflower do not require a specific vernalization

for transformation to the reproductive stage. Winter cauliflowers, and other biennial types,
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require cold treatment for curding and flowering, while for summer cauliflowers cold treatment

is only necessary for flowering. In the earliest group, i.e., high temperature, curd size is smaller
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than later groups. The curd color of Indian cauliflower is yellowish to cream white, while

Snowball is bright white. Nevertheless, variants having orange, green and purple color curds
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have been developed which are more nutritious than traditional white curds.
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An orange cauliflower mutant was first discovered in Bradford Marsh, Canada, in 1970.

Through many years of work by Prof. Emeritus Michael H. Dickson at Cornell University, and

other breeders, the orange cauliflower is now commercially available. The orange color results

from a novel spontaneous mutation of a single gene designated as Or for orange gene (Crisp et

al., 1975). The Or gene mutation causes the normally white curd to turn orange, and imparts

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visible orange coloration in the shoot meristems, the pith of the stem, and the vasculature at the

base of the petioles (Crisp et al., 1975; Dickson et al., 1988; Li et al., 2001; Howitt and Pogson,

2006). The Or homozygous plants (OrOr) have an intense orange coloration in these tissues,

exhibit stunted growth with a small curd and a delayed curd formation, while Or heterozygous

plants (Oror) are less pigmented and exhibit normal growth as wild-type plants, i.e., controlled

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by a single, semi-dominant, gene. The outer curd tissues of OrOr plants accumulate

approximately 8 mg·g-1 β-carotene, a level several hundred fold higher than that detected in

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comparable wild-type tissues (Li et al., 2001). The Or gene has been isolated by a map-based

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cloning strategy by Lu et al. (2006). It functions in increasing sink capacity/accumulation by

inducing formation of chromoplasts rather than altering expression of genes involved in

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carotenoid biosynthesis (Li et al., 2001; Lu et al., 2006; Lopez et al., 2008; Zhou et al., 2008).

‘Pusa Betakesari’, an orange curd cauliflower, contains 0.8-1.0 mg β-carotene in 100 g of curd
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tissue (Anonymous, 2016).

A purple cauliflower mutant displays vibrant violet hues resulting from a spontaneous
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mutation of a single, semi-dominant, gene designated Pr for purple gene. The purple coloration is
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due to accumulation of anthocyanins, a group of flavonoid compounds, which fulfill important

biological functions in protecting plants against biotic and abiotic stresses. They also exhibit
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antioxidant and anti-inflammatory activities, and may lower the risk for a number of chronic

diseases and cancer in humans. The mutant exhibited a tissue-specific pattern of anthocyanin
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accumulation, predominantly cyanidin 3-(coumaryl-caffeyl) glucoside-5-(malonyl)-glucoside.

Intense purple coloration occurs in the curd, young seedlings, very young leaves, very young

flower buds, siliques, and seed endosperm; while coloration was absent in older leaves, stems,

flower petals, and older siliques. The gene Pr encoded a R2R3 MYB transcription factor that

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exhibiting tissue-specific expression, consistent with an abnormal anthocyanin accumulation

pattern in the mutant. The curds accumulated approximately 3.75 mg·g-1 fresh weight cyanidin-

diglucoside (Chiu et al., 2010).

Breeding behavior

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The flowers of cauliflower are typically cruciferous, having 4 sepals, 4 petals, 6 stamens

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(2 are short) and 2 carpels. Two functional nectaries are present, situated between the bases of

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the ovary and short stamens, and the other 2 inactive nectaries are at the bases of pairs of long

stamens. Honey bees are the usual pollinating agents, though bumble bees and other syrphid flies

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also have roles in pollination. The stigma of Brassica species is receptive for 5 day before and 4
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day after anthesis, i.e., the longer receptivity is due to protogynous nature of the flowers. In

cauliflower, cross-pollination varies according to type, e.g., in summer cauliflower (‘Snowball’

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and ’Erfurt’), self-pollination may be as high as 70%; in winter and Indian cauliflower, cross-

pollination is high due to SI. The uses of SI, male sterility and doubled haploids are beneficial in
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commercial hybrid seed production.

Self-incompatibility
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Self-incompatibility is a mechanism that allows stigma to recognize and discriminate

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self-pollen preventing self-fertilization and inbreeding, and enforces out-crossing. Selfing could

be avoided by embryo abortion, but SI is pre-zygotic preventing embryo formation. Two types of
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SI have been reported (a) gametophytic and (b) sporophytic. In the sporophytic SI system,

incompatible pollen is manifested at the surface of stigma epidermal cells (papilla) by failure of

pollen grains to germinate and produce pollen tubes and is accompanied by deposition of callose

inside the papillae. There are 2 reasons for expression of SI: (i) lack of adhesion, hydration and

germination of the pollen grain, and (ii) failure to penetrate the papillae. Watts (1963) found

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higher SI in biennial winter and autumn types and lower SI in European summer types (Snowball

and Erfurt). Swarup and Chatterjee (1972) observed 90-100% SI among genotypes of Indian

cauliflower, while self-compatibility is dominant in the ‘Snowball’ group.

Homomorphic SI is generally controlled by a single S-locus containing 2 multiallelic

genes that encode the stigma-expressed S-locus glycoprotein (SLG), S-locus receptor kinase

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(SRK) and S-locus cysteine rich protein/S-locus protein 11 (SCR/SP11) described by Nasrallah

(2000) and Dickinson (2000). The allele forms of the locus are designated as ‘haplotypes’. The

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first 2, SLG and SRK, are determinants of SI specificity in the stigma. The SLG is a soluble cell-

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wall localized protein and SRK is a plasma membrane anchored signalling receptor, the extra

cellular domain of which shares similarity with SLG. The SRK gene is expressed in stigma
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papillar cells and encodes a plasma membrane-localized receptor kinase, which has a highly

polymorphic extracellular receptor domain (S domain) followed by a transmembrane domain and

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a serine/threonine kinase domain (Stein et al., 1991; Takasaki et al., 2000). However, SCR is the

male determinant, i.e., pollen coat-localized ligand, discovered by Nasrallah laboratory

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(Schopfer et al., 1999). It is a small, highly charged and polymorphic cysteine rich protein,

exclusively expressed in anthers during pollen development. The SLG gene, located at the S-
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locus, encodes a stigma soluble glycoprotein showing high similarity to the S-domain of SRK.
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Like SRK, SLG is a highly polymorphic protein between S-haplotypes. Because some S-

haplotypes lack the functional SLG gene at the S-locus (Suzuki et al., 2000), the SLG gene is not
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considered an essential component in SI in Brassica (Yamamoto and Nishio, 2014). The high

degree of identity shared by the 2 sequences suggests occurrence of gene conversion between

SLG and the S-domain of SRK in the same S-haplotype. This conclusion is supported by

analysis of progeny of a B. rapa commercial cultivars showing self-compatibility, in which part

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of the S-domain of SRK was replaced with the corresponding part of SLG from the same S-

haplotype (Fujimoto et al., 2006). Moreover, SCR is expressed in anthers and its translational

products secreted to the pollen coat (Iwano et al., 2003). It is a small peptide, ̴ 60 amino acids of

mature form, and functions as the ligand for SRK (Schopfer et al., 1999; Takayama et al., 2001).

SCR is highly polymorphic and less than 50% amino-acid sequence similarity is shared between

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S haplotypes (Schopfer and Nasrallah, 2000).

Annual Indian, and biennial winter, cauliflower have stronger SI mechanisms. A detailed

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investigation of SI in Indian cauliflower revealed that inbreds of maturity Group-I have the

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strongest SI followed by Group-II; Group-III had weak SI (Swarup and Chatterjee, 1972;

Chatterjee and Swarup, 1984; Singh et al., 2002). In self-pollination, stigma-localized SRK
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interacts with SCR of the same S-haplotype located on the pollen surface and activates a SI

signalling cascade, resulting in inhibition of self-pollen germination and pollen-tube penetration

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on the stigma papillar cell wall (Yamamoto and Nishio, 2014). A large number of S-haplotypes

have been identified in B. oleracea, B. rapa and in the closely-related R. sativus by pollination
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tests, electrophoretic analysis of stigma proteins, analysis of DNA polymorphism in SLGs or

SRKs, and determination of SLG, SRK and SCR sequences (Sakamoto et al., 1998; Oikawa et
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al., 2011). In B. oleracea, among the 49 S-haplotypes reported (Oikawa et al., 2011), sequences
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of the SLG, SRK or SCR alleles were determined for all S-haplotypes with the exception of the

S-haplotypes S-10, S-56, S-66 and S-67.

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The advantage of this system is the possibility to produce hybrid seed using 2 SI lines

homozygous for different S alleles as parental components. The disadvantages are less reliable

SI, which results in an undesirable number of “sibs” in hybrid seed and difficulties connected

with reproduction of SI lines through means of bud-pollination (BP). In cole crops, including

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cauliflower, it is necessary to maintain and multiply SI lines by breaking down the SI system.

Generally, SI lines are multiplied by BP, CO2 treatment and NaCl treatment.

Male sterility (MS)

Male sterility is the inability of the plant to produce fertile pollen, an efficient means for

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large-scale production of hybrid seed. It usually manifests in floral development as an

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incompatibility of nuclear-mitochondrial interaction in alloplasmic lines derived spontaneously.

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It may also arise following intraspecific, interspecific and intergeneric or wide crosses. In cole

crops, staminal MS systems are common. Stamenless male sterility (flower without stamens) in a

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natural population has been reported (Verma et al., 2014). Male sterility systems usually carry
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nuclear and mitochondrial genomes, i.e., genic male sterility (GMS) and cytoplasmic male

sterility (CMS), respectively. Use of biotechnological tools provides another type of male
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sterility, transgenic male sterility (TMS).

The GMS in cole crops are mainly recessive. A single recessive gene ‘ms’, mutated form
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male fertile ‘Ms’ gene, occurs in cauliflower (Cole, 1959; Nieuwhof, 1961). No marker genes are

linked to monogenic recessive male sterility, making it difficult to use this kind of male sterility.
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Van der Meer (1985) reported male sterility is under control of duplicate dominant genes with a

cumulative effect. Monogenic dominant male sterility has been described in cauliflower by
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Ruffio-Chable et al. (1993) and in other cole crops. This has limited possible practical value in
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hybrid seed production in some cauliflowers because of the unreliable nature of the SI system.

Some forms of GMS are temperature sensitive and result in self-pollination.

The CMS has not identified in cauliflower, or other cole crops, but has been introduced

from R. sativus, B. nigra, B. napus, B. juncea and B. tournefortii. CMS has been transferred in

broccoli through repeated backcrossing (Bannerot et al., 1974; McCollum, 1981). Hoser-Krauze

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(1987) transferred CMS from broccoli to cauliflower. The Ogura type CMS has been transferred

into heat tolerant Indian cauliflower from kale and broccoli, and is being used in heterosis

breeding. Pearson (1972) transferred CMS in cabbage from B. nigra, Chiang and Crete (1987)

introduced CMS from B. napus into cabbage and later from cabbage to cauliflower. The CMS is

maternally inherited, encoded in mitochondrial genes and can be utilized more effectively by

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breeders than GMS. Another set of nuclear genes, restorers of fertility (Rf), overcome the effect

of CMS genes, restoring the hermaphrodite condition which is cytoplasmic-genic male sterility

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(CGMS). Cytoplasmic male sterility can be considered a genetic system having 2 genetic

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determinants with different modes of inheritance. It involves variations at the cytoplasmic level,

with at least a sterility-inducing cytoplasm (S) and a cytoplasm with no sterility effect (N). At the
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nuclear level there is a dominant restorer allele (Rf) that enables plants with the S cytoplasm to

produce pollen and a recessive allele (rf), also called a maintainer of sterility, maintaining the
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male sterile phenotype induced by the S cytoplasm. The CMS system is an ideal solution for

hybrid development as crosses between a male sterile plant and a hermaphrodite plant
ed

(homozygous for maintainer alleles) give rise to 100% male sterile plants (Singh, 2015).

The most extensively investigated CMS system among cole crops started with the
pt

discovery of male sterility in Japanese radish (Ogura, 1968). After introgression through
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conventional breeding to B. oleracea, several developmental and floral abnormalities were

observed among CMS plants that affected their breeding value (McCollum, 1981). In B.
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oleracea, abnormalities of floral parts include petaloid and carpeloid stamens, petaloid anthers,

crooked style and reduced nectaries (McCollum, 1981). The Ogura and McCollum types of CMS

from radish exhibit chlorosis and loss of vigor when grown at lower temperature (<12°C) during

early stages of growth (Dickson, 1985). Improvement of female fertility and cold tolerance via

15
protoplast fusion (Pelletier et al., 1983; Jourdan et al., 1989) made Ogura CMS system, the most

popular one used for modern breeding of broccoli, cauliflower and cabbage F1 hybrids. Male

sterile cybrids with normal photosynthesis and improved nectar were obtained through

chloroplast exchange and mitochondrial recombination. The Ogura CMS system was first

improved in B. napus in 1983 and then in B. oleracea in 1989 (Delourme and Budar, 1999).

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Generally, the cytoplasm donor species, from which the sterility originates, provides the nuclear

restorers (Heyn, 1976; Delourme et al., 1995). The Ogura sterility in all B. oleracea species does

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not have any fertility-restorer genes, while all fertile cytoplasm forms act as maintainers (Prakash

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et al., 2009). In India, Ogura CMS is used to transfer the MS system in tropical and Snowball

cauliflower (Singh and Singh, 2016). The Ogura CMS lines of cauliflower, Ogu1A, Ogu2A,
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Ogu3A, Ogu12A, Ogu13A, Ogu14A, Ogu16A and Ogu33A have been developed (Dey et al.,

2011a, 2014).
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The first cyto-sterile B. oleracea plants with B. nigra cytoplasm developed by Pearson

(1972) through sexual hybridization were characterized by lack of nectaries and abnormal flower
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development (sepals, petals and styles). Low seed production ability with the B. nigra CMS

reported by breeders in cabbage, broccoli and cauliflower was a reason why this system was not
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used for practical breeding (Pelletier et al., 1983; Hoser-Krauze, 1987, 1989). Other practical
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problems are frequent presence of restorer genes (Rf) among B. oleracea populations having

nigra CMS background, and lack of known molecular/morphological markers useful for early
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identification of genotypes among fertile plants with the rf gene. Nevertheless, several male

sterile cauliflower plants with B. nigra cytoplasm have been developed through back-crossing

(Kaminski et al., 2012) with a higher ability for generative propagation had a similar

16
morphological structure in male fertile lines and did not show any deformation of styles, petals

and sepals.

The TMS is a biotechnology-based process developed to propagate seed of homozygous

male-sterile female inbred lines. One of the first biotechnology-based approaches to male

sterility, i.e., TMS was proposed by including tapetal-specific expression of a ribonuclease gene

t
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‘barnase’ to cause complete male sterility (Mariani et al., 1990). Tapetal-specific expression of a

ribonuclease-inhibitor gene ‘barstar’ in the male parent restores fertility to hybrid plants (Mariani

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et al., 1992). This technology is linked with the selectable marker phosphinothricin

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acetyltransferase conferring tolerance to the herbicidal active ingredient glufosinate ammonium.

Timely application of herbicide is essential to remove non-transgenic plants, which adds to cost
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and complexity. The specific expression of the cytotoxin can selectively destroy organs or tissues

related to pollen development, block the development process, and lead to male sterility. RNase
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digests RNAs. The genes encoding RNase— barnase and RNase T1 have been cloned. The

RNase gene and a specific promoter can be linked and transferred into plants to obtain male
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sterile plants. Mariani et al. (1992) fused TA29, the tapetum-specific promoter isolated from

tobacco anthers, with barnase gene from Bacillus amyloliquefaciens and with RNase T1 gene to
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be introduced into tobacco and oilseed rape through genetic transformation. They demonstrated
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introduction of male sterility in transgenic tobacco and B. napus plants. Genes coding for

ribonucleases were transferred independently, RNAase-T1 from Aspergillus oryzae and Barnase
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from Bacillus amyloliquefaciens. These genes, under control of the TA29 promoter 9 are

expressed in the tapetal cells. TMS has been developed in various crops, including cauliflower

(Janssens et al., 1992). There is no need to introduce the barstar gene in male parents of those

crops where sexual reproduction is not required for development of economic parts used.

17
Doubled haploid (DH)

In the classic breeding approach, production of F1 hybrids on a commercial scale is

largely based on inbreds that are selfed and selected through 6-8 generations of inbreeding,

which is tedious because of SI in B. oleracea crops. The DHs, produced by microspore culture,

t
are an ideal strategy to maintain parental lines because they generate inbred lines with 100%

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homozygosity only in 1 generation and are efficient in accelerating breeding, developing new

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varieties, doing basic genetic studies and saving time (Forster et al., 2007; Gu et al., 2014).

Microspore culture technology has been applied in Brassica breeding since the first report of

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successful isolation and culture of microspores in B. napus (Lichter, 1982) and in broccoli
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(Keller and Armstrong, 1983). There have been advances in microspore culture of different

Brassica species, especially cauliflower (Gu et al., 2014).

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Crop ideotypes

Most plant breeding is based on selection of superior, and/or elimination, of inferior

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traits. An additional approach is available through breeding of crop ideotypes, i.e., plants with

model characteristics known to require minimum resources, optimize morpho-physio-

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biochemical traits, and produce more yield. On the basis of genetic stocks available in the Indian
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cauliflower, it is possible to determine the most desirable ideotypes in cauliflower as having

specific maturity with 75-80% uniformity; 12-15 cm stalk length; plant type of No. 3 (self-
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blanch leaves); 40-50 cm frame size; 18-22 leaf number; 50-55 cm leaf length; hemispherical

curd shape; 15-18 cm curd diameter; 600-1000 g curd weight; 55-65 days of maturity; cremish-

white to white color curd; compact curd; devoid of riceyness, leafiness, fuzziness; and

resistance/tolerance to black rot, Alternaria blight, diamond back moth (DBM; Plutela

xylostella) and Spodoptera (Singh, 2015).

18
Breeding objectives

Crop varieties should possess the general features wide adaptability, superior uniformity,

ability to tolerate biotic and abiotic stresses, superior quality of produce, and high productivity.

The major breeding achievements for cauliflower are development of heat/humid tolerant

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cultivars, improved CMS system, and higher harvest index. Size uniformity is requested by the

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freeze processing industry. Smaller frame size, along with smaller curd, is becoming an

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important objective as it accommodates more plants per unit area and smaller curds are the

choice of nuclear/metro families. Curd colour is subject to consumer preferences and the aim is

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to obtain pure white, or creamy-white curds, or to develop colorful curds (orange, purple and
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green curds) with improved antioxidants, anthocyanins and carotenoid contents. Reduction of

glucosinolate content will allow a smoother taste, although the resultant cultivars are more
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sensitive to insect damage. The higher activity of the stress tolerance antioxidative enzymes

superoxide dismutase (SOD), catalase (CAT), guaiacol peroxidase (GPOX), ascorbate

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peroxidase (APX), glutathione peroxidase (GPX), glutathione reductase (GR),

monodehydroascorbate reductase (MDAR), dehydroascorbate reductase (DHAR) and

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glutathione S-transferase (GST) could play a role in improving tolerance to various type of

stresses. Efforts have been made to enhance antioxidant enzymes in vegetable crops through
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conventional breeding (Singh, 2007; Singh et al., 2009, 2010a, b). The most important
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cauliflower breeding objectives are:

i. high yield along with better crop ideotypes;

ii. uniform curding, i.e., uniformity in curd size, shape, maturity and color;

iii. compact and retentive white curds free from riceyness, leafiness and fuzziness;

19
 iv. high commercial quality as determined by size and shape, appearance, color persistence,

compactness and bruise resistance;.

v. small curds with smaller frame size, for dense transplanting;

vi. high harvest index, wide adaptability and better field standing ability;

vii. tolerant to heat/humidity, i.e., curd formation in summer/rainy seasons of North India;

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viii. robust SI and CMS lines to produce hybrids having better seeding ability; and

ix. cultivars tolerant to the diseases—black rot, Sclerotinia rot, downy mildew, Alternaria

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blight, Erwinia rot; insects—DBM, Spodoptera; and abiotic stresses—heat, rain, salt.

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Breeding methods an
During the 19th and 20th centuries, mass and recurrent selections were applied to achieve

greater uniformity and higher productivity. The F1 breeding started in the 1950s continue to
M
intensively harness the potential of heterosis to obtain harvest uniformity. In the 21st century, the

tools of molecular biology and biotechnology allowed important strides in hybrid breeding. The
ed

main breeding methods for cole crops, including cauliflower, are population improvement (mass

selection, stratified mass selection, gamete selection, family selection, line breeding, recurrent
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selection and mass-pedigree), combination breeding (backcross), utilization of the hybrid vigor

(synthetics, F1 hybrids), and utilization of molecular and biotechnological tools.

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Population manipulation has been used for improvement of cole crops. In India, mass
Ac

selection has been used for improvement of cauliflower. Though this method is useful for

improvement of simply inherited traits, it is not effective for traits governed by polygenes. Based

on progeny evaluation, modifications including mass-pedigree and family selection are better

than mass selection. The choice between these methods depends on the population, according to

the level of homogeneity due to SI. Family breeding is used to improve cauliflower and other

20
cole crops. In this method, seed of selected plants, based on progeny testing, are used to develop

synthetics. The disruptive selection method is recommended to break tight linkage in Brassica in

which extreme population types are selected and intermediate ones discarded.

Recurrent selection is a better option for improvement of quantitative characters,

especially those under control of additive gene action. This method has been effective for

t
ip
improvement of curd compactness, yield and other economic traits in cauliflower. Significant

improvement in yield (18-47%), and diameter, depth and weight of curd after 1 generation of

cr
recurrent selection has been reported (Tapsell, 1989). In India, variety Pusa Sharad, mid-

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maturity group, has been developed using recurrent selection (Sharma et al., 1999). The inbred

lines can be obtained by overcoming SI barriers, either with BP, or with chemical induction
an
methods (NaCl or CO2). The DHs are used to develop parental inbred lines. Use of inbreds as

cultivars is mostly applicable to summer European types which have self-compatibility. In this
M

approach, 2 self-compatible lines are crossed and the hybrid progeny is subjected to simple

pedigree/bulk/backcross method of breeding.

ed

Combination breeding has been used to transfer genes of interest through backcrossing.
pt

The donor parent is crossed to the recurrent parent followed by progenies which are then crossed

to the recurrent parent. The progeny of this cross is selected for the trait of interest and crossed
ce

back to the recurrent parent. This process is repeated for as many back crosses as needed to

create a line that is the recurrent parent with the gene of interest from the donor parent. The goal
Ac

of backcrossing is to obtain a line as identical as to recurrent parent with addition of a gene of

interest. This method was used to develop Pusa Shubhra, which is resistant to black rot, curd

blight and riceyness (Singh et al., 1993), and to develop CMS lines.

21
 Development of synthetic varieties is based on exploitation of additive genetic variance.

In a synthetic variety, unlike a variety developed by mass selection or line breeding, the desirable

lines having selected genotypes are synthesized after testing their general combining ability

(GCA). Synthetic varieties are produced in all Brassica vegetables after testing the inbred lines’

GCA. Generally, 4-8 lines are selected for developing synthetics. The advantage of synthetic

t
ip
varieties is: (i) its seed can be easily produced through open-pollination; (ii) it is particularly

useful where commercial seed industries are not developed; and (iii) it serves as a reservoir of

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germplasm. The cultivars Pusa Early Synthetic and Pusa Synthetic have been developed by

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synthesizing 6 and 7 parents, respectively (Singh et al., 1997; Gill, 1993).

Heterosis breeding is based on the principle of selecting parents, developing homozygous

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inbred/SI/CMS lines, testing combining ability, evaluating cross-combinations, and production

of commercial F1 seed. The F1 hybrids offer positivity, uniformity and consistency of vigor, yield
M

and quality, and are produced by crossing ≥2 inbred parental lines (Singh and Singh, 2016). In

cauliflower, heterosis was first reported by Jones (1932). At present, OP varieties of cauliflower
ed

have largely been substituted by F1 hybrids. Production of F1 hybrids is now developing faster,
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albeit with some technical difficulties, using improved Ogura CMS and DH parental lines.

Nieuwhof and Garretsen (1961) failed to find an appreciable amount of heterosis in European
ce

summer cauliflower (‘Snowball’, ‘Erfurt’, ‘Alpha’) which may be due to their narrow genetic

base. Watts (1965) and Swarup and Pal (1966) found significant amount of heterosis in
Ac

‘Snowball’ cauliflower. Swarup and Chatterjee (1972) investigated heterosis in Indian

cauliflower and reported better manifestation of it in the 1st maturity group>2nd maturity

group>3rd maturity group. Considerable amounts of heterosis have been reported by Singh et al.

(1975), Hoser-Krauze et al. (1982), Sharma et al. (2004), Kucera et al. (2006), Dey et al. (2011b)

22
and Jiafu et al. (2012). Heterosis for ascorbic acid, anthocyanin and carotenoids in cauliflower

has been reported by Dey et al. (2014). Heterosis for the stress enzymes SOD, POX and CAT has

been reported in cabbage (Singh et al., 2010b). Development of the robust system of genetic

emasculation approaches (SI, MS and DH) and inbred lines coupled with proper management of

pollinators will improve the economic aspects of seed production of F1 hybrids.

t
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Breeding for resistance to biotic stresses

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Black rot (Xantomonas campestris pv. campestris), Alternaria blight (Alternaria

brassicicola), downy mildew (Pernospora parasitica), watery soft rot (Sclerotinia sclerotiorum)

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club-root (Plasmodiophora brassicae), Verticillium wilt (Verticillium longisporum), Fusarium
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yellows (Fusarium oxysporum, F. conglutinanas), soft rot (Erwinia carotovora), turnip mosaic

virus (TuMV), powdery mildew (Erysiphae polygoni) and bottom rot (Rhizoctonia solanai) are
M
diseases affecting cultivation of vegetable Brassicas. Breeding for multiple disease resistance is

an important objective. Breeding lines have been developed with resistance to black rot, possibly
ed

the most serious disease of Brassicas worldwide. Sharma et al. (1972) evaluated parental lines

along with BC1, BC2 and F2 generations from 5 crosses involving MGS (highly resistant), Pua
pt

Kea and S. No. 445 (resistant), and S. No. 246, S. No. 15, EC 12013 and EC 12012 (susceptible-

highly susceptible); and suggested backcross breeding should be adopted, or to attempt selection
ce

in the progeny of a cross made between 2 resistant parents for evolving highly resistant lines.
Ac

Quiros and Farnham (2011) reported unstable inheritance of resistance of black rot because it is

influenced by the pathogen’s isolates. A segregation analysis of F2 progeny of a cross between

Pusa Himjyoti (susceptible) and BR-161 (resistant) indicated a single dominant locus governed

resistance to Xcc race 1, and observed 7 differentiating polymorphic markers (Saha et al., 2014).

23
 A relative resistance to several insect pests has been associated to a glossy leaf phenotype

(Quiros and Farnham, 2011). Trichome-based resistance against flea beetles was studied in B.

villosa (Palaniswamy and Bodnaryk, 1994) and resistance to the cabbage aphid (Brevicoryne

brassicae) was found in B. fruticulosa (Pink et al., 2003). Resistance to white fly (Aleyrodes

proletella) expressed by plants was found in B. villosa, B. incana and B. montana accessions

t
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(Pelgrom et al., 2015). In B. incana, presence of trichomes is likely responsible for observed

resistance to insects (Vosman et al., 2015).

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Transgenic

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Transgenic crops, commonly referred to as genetically modified crops, enable breeders to
an
bring favorable genes, often previously inaccessible, into already existing elite cultivars,

improving their value and offering unique opportunities for enhancing quality, modifying
M
morpho-phyisio-biochemical properties, and increasing tolerance to biotic (viral, bacterial

infections, pests and weeds) and abiotic (drought, frost, temperature, salt, UV rays) stress.
ed

Several strategies, including transgenic, have been used and developed to build resistance to

these stresses in plants. Among insect pests, the Lepidopteran larvae are the most problematic of
pt

vegetable Brassicas worldwide. The cry genes, cry1A, cry1Ab, cry1Ac, cry1A(b), cry1Ab3,

cry1Ba1, cry1C, cry1Ba1, cry1Ia3, and cry9Aa from Bt have been introduced into Brassica
ce

vegetables (Sharma, 1998; Zhao et al., 2006; Dias and Ortiz, 2014). Bt containing vegetable
Ac

Brassica successfully control the insect pests DBM, cabbage butterfly (Pieris brassicae), stem

borer (Hellula undalis), hairy caterpillar (Spilosoma obliqua) and cut worm (Spodoptera litura,

Agrotis ipsilon). In addition to cry Bt genes, the trypsin inhibitor genes of cowpea and soybean

confer resistance to Lepidopteran pests. Trypsin inhibitor, a protease inhibitor, is capable of

controlling a wide spectrum of insect pests (Zhao et al., 2006; Dias and Ortiz, 2012). Chitinase

24
genes cloned from plants and fungi have been transferred into vegetable Brassica that confer a

broad range of resistance to Alternaria blight, black rot and soft rot. In addition, various genes

are effective against fungal pathogens by employing glucanase to wire-stem, thionin to

Alternaria blight, permantis to damping-off and osmotin to white rust.

Various transformation approaches have been used in cauliflower, cabbage and Chinese

t
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cabbage to induce TMS. Toriyama et al. (1991) introduced an SLG gene from S8 homozygote of

B. rapa and were able to alter the SI phenotype of pollen and stigma. Transformations of the

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cauliflower plastid division gene, BoMinD have been reported using PEG-mediated

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transformation in mesophyll protoplasts (Nugent et al., 2006; Chikkala et al., 2013). Many

abiotic stresses lead to overproduction of reactive oxygen species which cause extensive cellular
an
damage and are major causes of plant damage as a result of environmental stress. Bhattacharya et

al. (2004) produced cabbage plants with enhanced salt tolerance through introduction of the
M

bacterial BetA, and Park et al. (2005) developed transgenic Chinese cabbage with the LEA gene

that allowed enhanced tolerance to drought and salt. The introduction of the choline oxidase
ed

(codA) gene from Arthrobacter globiformis into B. rapa L. spp. chinensis enhanced tolerance to

high temperature and high salinity due to accumulation of betaine (Wang et al., 2010). Tolerance
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to salinity was achieved by over-expressing the stress genes APX and SOD transgenically
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(Metwali et al., 2012). Further, Lu et al. (2006) developed transgenic cauliflower with high

levels of β-carotene accumulation. Wang et al. (2011) reported that RAPD, ISSR and SRAP are
Ac

the most powerful molecular markers for cultivar identification, genetic diversity analysis and

phylogeny study. The rapidly growing knowledge of the B. oleracea genome offers an

assembled sequence covering about 75% of the estimated genome size (Parkin et al., 2014), and

25
availability of thousands of markers and highly saturated genetic maps would certainly provide

increasing precision and extend options to facilitate breeding.

Future strategies

Strategies that have relevance to cauliflower breeding for enhancing production and

t
productivity, and efficient use of input resources are:

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i. population improvement to increase sustainability of production, enhance nutritional

cr
quality and reduce waste for which development of resistances to major abiotic and

biotic stress are recognised;

us
ii. efforts should be made to integrate 2- or multi-tiered breeding approaches for
an
broadening the genetic base, and introgressing genes and QTL of resistance, quality

and productivity into elite backgrounds;

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iii. difficulties in commercial seed production of F1 hybrids need to be overcome by

developing robust SI, Ogura CMS and DH parental lines, modern breeding techniques
ed

and proper management of pollinators, and

iv. rapidly growing knowledge of advance tools of parental line development, molecular
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markers and biotechnological interventions need enhanced utilization for improving

precision and extend options to support future breeding of cauliflower.

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us
an
M
ed
pt
ce
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41
Figure 1: U’s Triangle modified to include radish showing genome relationships among

cultivated Brassica species; solid and broken lines in allopolyploids represent female and

male parents, respectively. Genomes are represented by letters (A, B, C and R), and haploid

chromosome numbers are enclosed in parentheses, adapted from U (1935), Branca and

Cartea (2011), Prakash et al. (2011) and Singh (2015).

t
ip
cr
us
an
M
ed
pt
ce
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42
Table 1. Evolution of cultivated B. oleracea crops, adapted from Helm (1963), Prakash et
al. (2011), and Singh (2015).
Probable sequence of Scientific name
evolution (B. oleracea) Common name Ancestora
1 var. sylvestris L. Wild cabbage -
2 var. ramosa DC. Thousand-head kale, 1
branching bush kale
3 var. gemmifera DC. Brussels sprout 2
4 var. dalechampii 3

t
5 var. costata DC. Portuguese tree kale, 1

ip
tronchuda kale
6 var. medullosa Thell. Marrow-stem kale 1

cr
7 Intermediate between 6 and 8 6
8 var. gongylodes L. Kohlrabi 7
9 var. sabauda L. Savoy cabbage 5

us
10 var. capitata L. White cabbage 9
11 var. capitata L. Red cabbage 10
12 var. viridis L., var. sabellica L., Kale and collards 1
var. palmifolia DC.
an
13 var. italica Plencks Broccoli, Calabrase 12
14 var. botrytis L. Cauliflower (biennial) 13
15 var. botrytis L. Cauliflower (annual) 14
16 var. botrytis L. Cauliflower (Indian) or 15
M
Tropical cauliflower
a
Ancestors represented by numerical letters (1 to 15) of order of evolution along with
corresponding scientific name.
ed
pt
ce
Ac

43
Table 2. Nomenclature of cultivated B. oleracea L., adapted from Anonymous (2015) and
Maggioni (2015).
Scientific name
(B. oleracea var.) Common name Common synonym
botrytis L. Cauliflower
capitata L. Red/White/Shetland cabbage
sabauda L. Savoy cabbage
gemmifera DC. Brussels sprout
gongylodes L. Kohlrabi B. caulorapa (DC.)

t
Pasq.

ip
italica Plencks Broccoli
viridis L. Kale, collard var. acephala DC.

cr
medullosa Thell. Marrow-stem kale
ramosa DC. Thousand-head kale, subsp. fruticosa Metzg.
branchingbush kale

us
sabellica L. Curly kale
palmifolia DC. Palm kale, Jersey kale
costata DC. Portuguese tree kale, tronchuda
kale
an
alboglabra (L.H. Bailey) Musil Chinese kale, Kalian B. alboglabra L.H.
Bailey
M
ed
pt
ce
Ac

44
Table 3. Classification of cauliflower grown in India on the basis of maturity period and
temperature requirement, and recommended varieties for cultivation.
Maturity OTRCIDa
group Period of maturity (°C) Variety/Cultivar/Genotype
Ia (Early) September Kunwari 20-27 Punjab Kunwari, Early Kunwari, Pusa Early
Synthetic, Pusa Kartik Sankar, Pusa Meghna,
Kashi Kunwari
Ib (Early) October Katki 20-25 Pusa Deepali, Pusa Katki, Pant Shubhra, Pant
Gobhi-3, VRCF-86

t
II (Mid) November Agahani 16-20 Improved Japanese, Pant Gobhi-2, Pant Gobhi-4,

ip
Pusa Hybrid-2, Pusa Hybrid-3, Pusa Sharad,
Punjab Giant-26, Pusa Agahani, Kashi Agahani,

cr
VRCF-50, VRCF-102
III (Mid late) December Poosi 12-16 Pusa Synthetic, Pusa Shubhra, Pusa Himjyoti,
Punjab Giant-35, Hissar-1, D-96, VRCF-22,

us
VRCF-202
IV (Late) January Maghi 10-16 PSBK-1, PSB-1, PSB-2, Dania, Ooty-1, Kala
Patta, Pusa Paushja, Pusa Shukti
a
OTRCID = Optimum temperature range for curd initiation and development.
an
M
ed
pt
ce
Ac

45
Table 4. Global germplasm collection of B. oleracea species at various gene-banks, adapted
from Anonymous (2010).
Accession Type of accession (%)
a
Gene-bank Number % share WS LR BL AC OT
Science and Advice for Scottish 2367 11.7 1 99
Agriculture, Edinburgh, UK
Northeast Regional Plant Introduction 1625 8.1 6 1 5 88
Station, Plant Genetic Resources Unit,
United States Department of

t
Agriculture, Agricultural Research

ip
Services, New York State Agricultural
Experiment Station, Cornell University,

cr
Geneva, NY
Beijing Vegetable Research Centre, 1235 6.1 100
Beijing, China

us
Genebank, Leibniz Institute of Plant 1215 6.0 2 32 3 60 3
Genetics and Crop Plant Research
(IPK), Germany.
Unité Expérimentale de Sophia- 1200 5.9 100
an
Antipolis, Groupe d’Étude et de
Sophia-Antiopolis contrôle des
Variétés et des Semences, France.
NI Vavilov All-Russian Scientific 980 4.9 26 74
M
Research Institute of Plant Industry,
Russian Federation.
National Institute of Agrobiological 672 3.3 1 7 91
Sciences, Japan.
ed

Centre for Genetic Resources, the 631 3.1 12 2 75 11

Netherlands.
Others (98) 10257 50.8 3 24 5 34 35
pt

Total 20182 100 2 16 3 33 46

a
WS = wild species; LR = landraces/old cultivars; BL = breeding lines; AC = advanced
cultivars;
ce

OT = others (the type is unknown or a mixture of 2 or more types).

Ac

46