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Breeding of Ornamentals: Gladiolus

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Breeding of Ornamentals: Gladiolus

S.K. Datta
Retired Scientist, CSIR-National Botanical Research Institute, Lucknow, Uttar Pradesh, India
Ex CSIR Emeritus Scientist, Bose Institute, Kolkata, West Bengal, India
Email id: subodhskdatta@rediffmail.com

Received: 21-02-2020; Accepted: 18-05-2020

ABSTRACT

Gladiolus is grown all over the world for cut flower production for floriculture trade. Extensive
research has already been done on gladiolus to generate basic knowledge on multidisciplinary
aspects. Efforts in germplasm collections, characterisation, cytology, tissue culture, disease
management, classical improvement and induced mutation are worth mentioning. Despite significant
technological advances, we are still repeating some of our experiments as routine activities. Selective
breeding is now most essential to develop varieties as per requirement of the trade. This will be
possible in two ways-utilising all available knowledge as guidelines and identification and wise
selection of parents having desirable genes prior to cross-breeding.

Keywords: Gladiolus, Germplasm, Genetic diversity, Hybridization, Pigments, Mutation,

Biotechnology

INTRODUCTION and delicate florets of various shades, sequential opening

of flowers for a longer duration, good keeping quality of
The floriculture sector is expanding rapidly all over the cut, ease of growing and multiplication and suitability for
world, including India. Floriculture is now acknowledged open cultivation (Singh, 2006). The number of literature
on the whole as profitable occupation. Ornamental crops published on gladiolus, worldwide, in general and India in
with their characteristics and unique admirable qualities particular, is so voluminous that it is difficult to highlight
are widely cultivated worldwide. Natural diversity of the details of each publication. But there are no specific
ornamental plants plays diverse role in economic terms publications that highlight the entire research activities on
for human society. Gladiolus is now the most important different aspects of gladiolus. The time has come to assess
cut flower in floriculture industry. Gladiolus is one of the the present status of multidisciplinary research on gladiolus
world’s most popular flower due to its long history, colour and its impact on floriculture industry. The journey of
and sheer elegance of form. Gladiolus is cultivated research on gladiolus from its starting point to its present
commercially worldwide where spring and summer scenario has been illustrated in the present article. Books,
conditions are favourable. Scientific and commercial internet sources, relevant reports of researchers of various
advancements of gladiolus are noteworthy due to institutions and universities and author’s own experience
increasing demand in trade and interdisciplinary scientific have been used as resource to prepare the article. The
excellence. It has become one of the lucrative cut flowers article focuses the early history and practical breeding
due to its majestic attractive spike forms and size, elegant and related research on gladiolus. Extensive research has

Volume 9, Number 2, May-August, 2020, pp. 115-133 115

 S.K. Datta

already been done to generate basic knowledge on successful experiments. The author tried to gather the
multidisciplinary aspects of gladiolus. Now, there is a need information scattered worldwide and in India in a sort of
to formulate future research programme keeping in view summary or general agreement of what is known on
the changing priorities and thrust areas of floriculture. It breeding at the moment. For optimum utilisation of any
is high time to analyse the research activities on gladiolus commercial crop proper understanding of the evolutionary
in our country. There is a need for proper designing of dynamics, elemental ecogeographical distribution, species
breeding research on a need basis. The article presents a and cultivar range, knowledge of its breeding system,
total scenario on diverse features of research for agro-technology, techno-economics, genetic variability
improvement of gladiolus. This may help to redesign the etc. are essential. The present write-up is an attempt to
future research plan to get the desired results quickly as unfold the journey of gladiolus research from its starting
per desire in floriculture. It is not possible to analyse all point to its present scenario. To develop a conceptual
research topics and related publications. The salient picture of the theme it is essential to begin from the early
features of all major activities have been highlighted with research initiative and gradual development.
restricted reference. One can see all publications from
cited references and cross references. Though meticulous ORIGIN AND GEOGRAPHICAL DISTRIBUTION,
care has been taken in reviewing but while dealing with GENUS, SPECIES AND CULTIVARS
such a voluminous work, some mistakes/errors might
have crept in, despite my best efforts. Since the days of ancient Greece, the gladioli are said to
be cultivated. The gladiolus cultivation in Brazil started in
In spite of publications of huge literature, there is not a the years 1950s (Tombolato et al., 2002). It is known
single source where all research developments on gladiolus since 1578, as evidenced by a record in Lyte’s Nievve
are documented. There is diverse opinion about its Herball, first were introduced into France. The gladiolus
distribution, total number of species and varieties, is produced in Australia, United States, Canada, Italy,
characterisation and improvement. Nearly all present France, Holland, India, Poland, South Africa, England and
research approaches are independent, repetitious and Japan, among others (de Hertogh and Le Nard, 1993).
beginning from starting point. There are few publications South Africa, and particularly the Cape Province, is
that highlight the research activities in India on different considered the centre of diversity of the genus. Two
aspects and components of floriculture. Considering the species are found in Madagascar and fifteen in the
increasing business potential of floriculture, it is high time Mediterranean area (Delpierre and Plessis, 1974; Lewis
to judge the present status of research and its impact on et al., 1972; Ohri and Khoshoo, 1985a). Species of
floriculture industry. Recently, author has taken careful Gladiolus occur in Africa and Madagascar, in Europe and
efforts to review the floriculture research (Datta, 2019a, western Asia (Hamilton, 1980). The majority of the ~260
2019b) and its different components like Chrysanthemum Gladiolus species originate from South Africa and the
(Datta, 2013; Datta and Janakiram, 2015), Rose (Datta, Cape of Good Hope is considered to be the centre of
2018) and Tuberose (Datta, 2017a). It is high time to diversity of this genus (Goldblatt, 1996; Valente et al.,
evaluate the complete research activities conducted on 2011). The second centre of this species richness is in
gladiolus in our country. Considering the important the Mediterranean region with only about seven species
position of gladiolus in Indian floriculture, voluminous originating from this region (Valente et al., 2011). Almost
research has been conducted on this crop on all of these grow in sub-Saharan Africa (Goldblatt et al.,
standardisation of agro-techniques, utilisation of agro- 2001). These were the only wild species as garden gladioli
chemicals, development of new and improved varieties, three hundred years ago (Hartline, 1996; Cantor et al.,
post-harvest, breeding system, disease management, etc. 2010). The present days Gladiolus cultivars are complex
at different research institutions and universities. A large hybrids and include the following species: Gladiolus
number of publications are available as a result of cardinalis Curtis., G. dalenii van Geel., G. oppositiflorus

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 Breeding of Ornamentals: Gladiolus

Herb., G. papilio Hook. f., G. carneus Delaroche, G. 1989). Majority species did not participate in the
cruentus Moore., G. tristis L., and G. saunddersii Hook. development of modern cultivars but may possess useful
f. (Huxley et al., 1992). Modern gladioli are generally character, for plant breeding. 227 species have been
grown as summer-growing cut flowers which are reported reported by Arora et al. (2002). The wild species of gladioli
to have derived from summer-growing species (G. are native to the Mediterranean region, the Middle East,
dalenii, G. oppositiflorus, G. papilio) and winter growing western Asia, Madagascar and South Africa. 10,000
species (G. saundersii, G. cardinalis) through named cultivars were probably derived from just seven
hybridization. Survey of early and recent literature does species native to South Africa (Kispotta et al.,
not give a clear picture. Reticulate evolution of present- 2017a,2017b). It contains 180 species with ploidy level
day decorative varieties by means of ongoing hybridization, ranging from 2n=2x=30 to 2n=12x=180 (Larson, 2012).
introgression cycles and selection have played an important All the modern cultivars were developed from few wild
role in the diversification and evolution of Gladiolus L. species and are considered as tetraploids (Ohri, 2013;
(Barnard, 1972; Ohri and Khoshoo, 1983a, 1983b; van Manzoor et al., 2018). Its origin is South Africa and
Raamsdonk and de Vries, 1989; Goldblatt et al., 2001). It comprises of about 300 species in which 250 are wild
is a matter of debatable question about the exact number and 50 of garden origin (Kispotta et al., 2017a,2017b).
of species under the genus Gladiolus. Gladiolus is The present cultivated gladiolus have been developed from
represented by about 180–300 species. Most of the 180 20 to 25 species and is cultivated in almost all countries
Gladiolus spp. are reported from South Africa (Duncan, of the world where spring and summer conditions are
1982; Goldblatt, 1996; Goldblatt et al., 1993), although favourable (Cantor and Tolety, 2011; Kumari et al., 2016).
they are widely distributed to as far north as Russia and Modern Gladiolus x hybridus developed from 20 to 25
into the Mediterranean. 72 species have been identified as species. In the Cape Province region alone, as many as
being native in the Cape Province region (Kidd, 1996). 72 species have been identified as being native (Kidd,
The number of species in the genus is 255 and many of 1996; Anderson et al., 2012). Gladiolus cultivation in India
them are found in South Africa, Tropical Africa, dates back to the 19th century as ‘Firmingers Manual of
Madagascar and Eurasia (Goldblatt and Manning, 1998). Gardening in India’ published in 1863, mentions that
Present gladiolus is considered to have been bred originally Charles Gray of Coonoor grew some gladioli from corms
from only six species (Lewis et al., 1972; Shaukat et al., and seeds in his garden. The major states growing gladiolus
2013). This genus is represented by about 250 to 300 are Uttar Pradesh, Himachal Pradesh, Haryana, Delhi,
species (Tombolato et al., 2002). The number of species Karnataka, Maharashtra, Punjab, West Bengal, Odisha,
in the genus is 255 (Pragya et al., 2010a) and more than Chhattisgarh, Assam, Sikkim and Meghalaya, Uttarakhand
10,000 cultivars of which 20 are grown commercially (Naresh et al., 2015; Verty et al., 2017; Pattanaik et al.,
for cut flower purpose (Naresh et al., 2015). All 300 2015). Thus gladiolus is considered as winter crop in
species of the genus grow in sub-Saharan Africa (Goldblatt plain areas and summer crop in hilly areas (Aftabet al.,
et al., 2001; Cantor et al., 2010). The number of species 2007). Gladiolus was introduced in Bangladesh around
in the genus currently totals 255 (Goldblatt and Manning, 1992 from India (Mollah et al., 2002; Hossain et al., 2011).
1998; Kumari et al., 2016; Takatsu et al., 2002). More
than 255 species are reported in the genus that grows in Classification
western and central Europe, western Asia, and in tropical
and southern Africa (Cantor et al., 2011). Rashmi and Gladiolus has been classified as follows: Botanist grouped
Kumar (2014) reported 260 species. Goldblatt and gladiolus species into four groups on the basis of their
Manning (1998) reported 255 species and present day geographical origin—(1) Eurasian group (G. atroviolaceus,
cultivars of Gladiolus x grandiflora developed from a G. communis, G. cardinalis, G. floribundus sp. Miniatus,
small number of wild species G. cruentus, G. natalensis, G. floribundus subsp. Rudis, G. palustres), (2) East
G. oppositiflorus, G. papilio and G. saundersii (Imanishi, African group (G. aequinoctalis), (3) Natalensis group

Volume 9, Number 2, May-August, 2020 117

 S.K. Datta

(G. psittacinus var. hookeri, G. psittacinus var. cooperi, Unfortunately, there is no correct system for authentication
G. psittacinus var. dracocephalus, G. primulinus) and (4) of each variety. Number of varieties available in different
South African Cape species (G. maculates, G. odoratus, centres in India has been reported by different authors
G. oppositiflorus, G. orchidiflorus, G. saundersii, G. from time to time. ICAR under All India Coordinated
sempervirens, G. tristis; Parthasarathy and Nagaraju, Research Project on Floriculture took initiative in 2002 to
1999). Gladiolus has also been classified into six groups compile passport data of all the ornamental varieties
on the basis of plant height, flower size and their including gladiolus (Bhattacharjee et al., 2002). Total
arrangement on the spike: Grandiflorus or large-flowered number of gladiolus varieties reported from each centre
hybrids, Primulinus hybrids, Butterfly hybrids, Miniature for passport data is as follows—Maharana Pratap
hybrids Face ups, Colvillei hybrids, Ochideola (Arora et University of Agriculture and Technology, Udaipur = 63;
al., 2002). North American Gladiolus Council (NAGC) PAU, Ludhiana = 36; G.B. Pant University of Agriculture
has recommended a broad system of classification of and Technology, Pantnagar = 42; Sher-e-Kashmir
gladiolus based on floral characters. It has been classified University of Agricultural Sciences and Technology,
into three groups on the basis of plant height (cm)—Tall Srinagar = 60; Mahatma Phule Krishi Vidyapeeth, Pune =
(>100), Medium (50–100) and Dwarf (up to 50). 50; Acharya N.G. Ranga Agricultural University,
Gladiolus varieties are also classified according to flower Hyderabad = 27; Birsa Agricultural University, Ranchi =
size—diameter (cm)—Giant gladiolus (>14), Large 15; IARI, New Delhi = 47; IIHR, Hessaraghatta = 74;
gladiolus (11.4–14), Medium gladiolus (8.9–11.4), Small Indian Agricultural Research Institute Regional Station,
gladiolus (6.4–8.9) and Miniature gladiolus (<6.4). Katrain = 29. CSIR-NBRI Lucknow reported total
Cultivars are classified on the basis of blooming time germplasm 105. But, there are duplications. Name of
(days) from the date of planting the corms—Early Season varieties maintained at different centres, varieties are
(within 60), Mid-Season (60–90) and Late Season (>90; grown extensively in many parts of the world, varieties
Sharga and Roy, 2002). for cut flowers in world trade, Indian bred varieties, etc.
have been mentioned in different publications (Datta, 2001;
Varieties Reported from India Arora et al., 2002; Sharga and Roy, 2002; Bhattacharjee
et al., 2002).
It is imperceptibly difficult to mention the exact number
of total varieties available throughout the world. Different Genetic Diversity
workers have reported the names of varieties available
with them. There are duplications in names of varieties. Genetic variability is a basic pre requisite for plant breeding
Gladiolus research in India is going on in some research programme on which selection acts to evolve superior
institutions, Universities and the major activities are genotype. The higher the amount of variation for different
coordinated by DFR through its network at different characters in the chosen material, the greater is the scope
coordinating centres (Udaipur; IIHR Hessaraghatta; BCKV for its improvement through selection. A wide range of
Kalyani; Srinagar; PAU Ludhiana; Y.S.P.U.H.F. Solan; variation is generated when genetically diverse and
Hyderabad; IARI New Delhi; Pusa Bihar; Pune; Pashighat; geographically isolated lines are brought together. The kind
Ooty and CSIR-NBRI). The number of total germplasm/ of genetic diversity detected among the genotypes
varieties is incredibly large in India as per reports from belonging to same geographical region might be due to
different centres. But it is very difficult to say the exact differences in adaptation, selection criteria, selection
number as each centre collects varieties from other centres pressure and environmental conditions. Variability
through sale/exchange/research purpose/trial, etc. information about the different characters present among
Therefore, there is duplication of varieties which reflects the individual cultivars belonging to single species or
on total germplasm collection. In addition, same varieties different species is important for the future breeding
are named in different names at different places. programme. It also helps to access the extent of combining

118 LS - An International Journal of Life Sciences

 Breeding of Ornamentals: Gladiolus

ability between two different contributing characteristics. taken for spike emergence was negatively and significantly
Exploitation of variability is of great importance and is correlated with number of floret per spike, number of
prerequisite for the effective screening of superior shoot per plant and vase life. High heritability and high
genotype (Kispotta et al., 2017a,2017b). The heredity and genetic advance in spike length indicate the presence of
variation of any taxa are controlled by its genetic system. additive gene effects in these traits and their amicability
This knowledge helps to prepare a meaningful breeding for direct selection (Kispotta et al., 2017a,2017b).
methodology for genetic improvement (Ornduff, 1969). Genotypic correlation coefficients were higher than the
The extent of genetic variability, heritability, genetic phenotypic correlation (Balaram and Janakiram, 2005;
advance, correlation and path coefficient of a large number Kumar et al., 2012). Significant positive association of
of available gladiolus genotypes were studied by different number of florets per spike was observed with spike
scientists and reported from time to time and recently length (Monika et al., 2008; Choudhary et al., 2011;
maximum work is going on in this direction. All Lahijie, 2012; Pal and Singh, 2012; Bhatia and Grewal,
experiments were designed to study the phenotypic co- 2009; Jhon et al., 2002; De and Misra, 1994; Patra and
efficient of variance (PCV) and genotypic co-efficient of Mohanty, 2015). High PCV and GCV for rachis length
variance (GCV) for almost all the characters of gladiolus. and low for number of spike, floret diameter, floret length
Considerable amount of variation was observed for all and number of leaves, leaf width, plant height, spike length,
the characters under study. The genotypic and phenotypic spike weight, weight of daughter corm and number of
coefficients of variation are required to understand the cormels per corm, corm weight per plant, were observed
effect of environment on various polygenic traits. The (Anuradha and Gowda, 1990; Balaram et al., 2000; Geeta
degree of inter relationship among various characters and et al., 2014; Rashmi and Kumar, 2014). More variability
in evolving selection criteria for improvement of the in corm weight and spike weight is indicated by high value
available germplasm correlations between different of PCV along with GCV. High GCV noticed for number
characters are very helpful. Genotypic correlation was of side shoots, number of cormels per plant, plant height,
of higher magnitude than phenotypic correlation in most number of florets per spike and days to flowering
of the cases. The results of correlation coefficient reveal (Balamurugan and Arijmugam, 2002; Bichoo et al., 2002;
that for yield improvement through selection, much Pratap and Manohar, 2006). Low GCV observed for
emphasis should be given on the characters like spike longevity of individual florets and duration of first floret
yield, corm yield, plant height, spike length, days to spike (Balamurugan and Arijmugam, 2002). Significant positive
initiation, days to floret initiation, number of florets, corm correlation between plant height with spike length has
weight, spike weight, corm diameter, etc. (Pattanaik et been reported (Gowda, 1989; Neeraj and Jha, 2001). Misra
al., 2015; Soorianath Sundaram and Nambisan, 1991; and Saini (1990) reported positive significant correlation
Maitra and Satya, 2004). Corms weight per plant was of number of florets per spike with height of the plant.
significantly and positively correlated with spike weight, Deshraj and Mishra (1998), Neeraj and Jha (2001) reported
number of cormels per plant, number of corms per plant, significant positive association of number of florets per
spike initiation, number of floret per spike and number of spike with plant height. Maitra and Satya (2004) reported
leaves per plant (Rashmi and Kumar, 2014). that days to flower bud initiation exhibited high negative
correlation with plant height, spike length, number of
Highest PCV and GCV coefficients of variation were florets spike-1. Naresh et al. (2015) studied eight hybrids
observed for the spike length followed by number of floret for 20 characters the genetic variability, heritability, genetic
per spike, days taken for the spike emergence was advance and correlation for identifying suitable strains
positively and highly significant with the days taken for for coastal conditions of Andhra Pradesh, India. The
bud initiation, days taken for 1st floret to show colour, magnitude of phenotypic coefficient of variation (PCV)
days taken for 1st floret to open, number of floret open was higher than genotypic coefficient of variation (GCV).
at a time, diameter of the floret and spike length. Days Heritability used in conjunction with genetic advance

Volume 9, Number 2, May-August, 2020 119

 S.K. Datta

provides better information for selecting the best should have clear breeding objectives. For a meaningful
individuals than the heritability alone. Grafius (1965) breeding project collection of sufficient gene pool and
mentioned that the optimal genetic level for each identification of desirable genes/genotypes are very
component would differ depending on the type of the important. This will help in selective crossing to develop
environment. The results of maximum correlation more favourable combinations. First of all the breeders
coefficient investigations indicate that spike yield and corm should collect germplasm and to identify the valuable
yield are the two most important characters which should accessions through characterisation. Another aspect of
be considered for yield improvement of gladiolus the breeding is to collect wild species and old cultivated
population. Hence, these characters may be considered varieties and to use new exotic species to create new
as selection indices in gladiolus breeding programme. products. In gladiolus, there is a continuous development
of new varieties through breeding and sufficient
Cultural Practices knowledge on all the above aspects that have been
generated. But unfortunately, the background information
Considering the important position of gladiolus in Indian regarding practical approach for success of breeding has
floriculture, sufficient quantum of work and research not been adequately documented and also documented
results/methodologies developed at various stages of information have not been properly utilised. In this article,
research have been reported from time to time on different author tried to provide a synthesis of relevant information
components. The main area of early and subsequent on the subject. Important features of gladiolus breeding
research in India concentrated on following aspects— in the context of results reported by different breeders
standardisation of agro-techniques (under wide agro- and based on author’s experience are discussed. No
climatic conditions including sodic wasteland), utilisation technology has been discussed in detail, only the achieved
of agro-chemicals (effect of nitrogen, phosphorus, potash, knowledge and few interesting examples of specific
cadmium, etc.), foliar application (urea),role of corm size, varieties have been cited as ready reference. To develop a
spacing, time of planting, corm and cormel production, conceptual picture of the theme it is essential to begin
techno-economics, development of new and improved from the beginning of early breeding history and gradual
varieties, post-harvest (effect of seasonal harvest, development. The history of early breeding ideas will
packaging, micronutrients, temperature, effect of cobalt, develop insight about actual changes in approach in
silver thiosulphate, aluminium sulphate, ethyl alcohol, development of recent breeding research both
sucrose, citric acid, AlSO4, 8-HQC, NAOCl, Ca(NO2), quantitatively and qualitatively. The expected results may
CoSO4, STS, NiCl, Al2(SO4)), breeding system, disease be achieved more easily and in less time if the past
management, etc. at different research institutions and experiences are used properly.
universities. Huge amount of publications are available as
a result of successful experiments (Arora et al., 2002; Factors of Selection
Datta, 2001; Sharga and Roy, 2002; Mukhopadhyay, 1995;
Bose and Yadav, 1989). The main objectives of gladiolus breeding generated from
different scientists are—to develop varieties with
Breeding agronomic characteristics of high productivity, high rate
of germination of the cormlets, vigorous corms which
Many floricultural crops are vegetatively propagated and grow under several conditions of soil and climate, early
are highly heterozygous, which causes a complex flowering, attractive/novel floret colours and floret
inheritance of genetic factors as well as the polyploidy, arrangement on the spikes, large sized florets, long spike,
making it difficult to improve through conventional increased number of florets, good stem strength, prolonged
breeding (Shibata, 2008). Breeding is the purposeful flowering period, resistance to major pest and diseases,
manipulation in order to create desired genotypes and long vase life, flowers with high ornamental and
phenotypes for specific purposes. Therefore, breeders commercial qualities, resistance to the great variation of

120 LS - An International Journal of Life Sciences

 Breeding of Ornamentals: Gladiolus

temperature and luminosity, flowers easy to manipulate, species, including hardy ones derived from G. angustus,
pack, store and capacity to flower after 2–3 days of dry G. cardinalis, G. carneus and G. tristis, and tender winter-
storage, good corm multiplication rate, winter flowering growing ones. Others also started breeding Gladiolus,
capability, flower fragrance, etc. including Colville’s of Chelsea, who crossed G. cardinalis
and G. tristis to make G. x colvillei, described in 1826
Heritable traits such as yield and quality are complex and still cultivated (Chis et al., 2010). In 1874, Max
characters known to be collectively influenced by various Leichtlin obtained the first gladiolus hybrid from the cross
polygenically inherited traits that are highly susceptible to of G. gandavensis Van Houtte with G. saundersii Hook.
environmental changes. Therefore, the knowledge of
The modern hybrids are designated as G. X grandiflorus,
direction and magnitude of association between yield and
which is a complex of about 11 species like G. carneus,
quality and their components and within components
G. dalenii, G. oppositiflorus, G. gandavensis, G.
themselves become necessary for an effective and
primulinus, G. papilio, G. saundersii, etc. In varietal
efficient selection of genotypes in gladiolus for yield and
assessment experiments a number of varieties have been
quality parameters. Gladiolus is a very suitable material
identified which are less susceptible to the day length,
for breeding due to its heterozygous nature and polyploidy
more resistant to the low temperature, produce long
status. The breeding program is an important
spikes, more adapted to the winter production, well
preoccupation of many researchers in the whole world.
adapted to summer climate, etc.(Anderson and Park,
It is constant work to introduce permanently new cultivars
1989; Ohri and Khoshoo, 1985b; Tombolato et al., 2002).
to increase the variability of the Gladiolus species (Hort
New varieties have been developed with extended vase
et al., 2012). The present days Gladiolus cultivars are
life, floral novelty, extended flowering periods, etc.
complex hybrids and following species are mostly
(Takatsu et al., 2002). Winter-hardy species from Russia
involved in breeding: G. cardinalis Curtis., G. dalenii van
include G. imbricatus and G. palustris. In addition, several
Geel., G. oppositiflorus Herb., G. papilio Hook. f., G.
species are adaptable to cultivation, including G. alatus,
carneus Delaroche, G. cruentus Moore., G. tristis L., and
G. angustus, G. cardinalis, G. carmineus, G. carneus, G.
G. saunddersii Hook. f. Hybridization between wild species
and cultivars of Gladiolus is an effective means of dalenii, G. ochroleucus, G. pritzelii, G. saundersii, and G.
producing individuals (genotypes) with desirable features sempervirens (Duncan, 1982). Modern gladioli are
(Ohri and Khoshoo, 1983b; Cantor and Tolety, 2011). primarily grown as summer-growing cut flowers and
However, Eurasian species have not been used in tender annuals. They are derived from summer-growing
developing modern cultivars of gladiolus even though they species, including G. dalenii, G. oppositiflorus, G. papilio,
can be valuable because of their relative hardiness and and G. saundersii. Gladiolus cardinalis, a winter growing
low sensitivity to fungal diseases (Ohri and Khoshoo, species (winter rainfall region), has also been used in
1983b; Rakosy-Tican et al., 2012). Hybridization between hybridization. In South Korea breeding work started in
closely related species (G. imbricatus, G. italicus and G. 1986 and the first reddish cultivar (‘Hongkwang’) and
illyricus) has occurred as a result of migrations due to the bright orange-based (‘Hongeun’) were released in
climatic changes (van Raamsdonk and de Vries, 1989). 1995. More than 49 new cultivars were released from
Moreover, G. communis subsp. byzantinus appears to 1995 to 2008 comprising new colours, very early and
hybridize with G. illyricus within its core range in southern very late flowering, resistance to thrips (‘Pink Smile’),
Spain producing an evenly-graded range of morphological etc. (c.f. DH Goo, HY Joung, YI Kang, YJ Choi, HK Shin,
intermediates (Cantor and Tolety, 2011). Low seed set in Gladiolus Breeding in Korea). Desirable qualities (flower
a few intervarietal crosses and failure of seed set in some form and shape, colour diversity or scent, etc.) of
interspecific crosses are apparently realised as major individuals might have produced through hybridization
problems in breeding of gladiolus (Van Tuyl, 1997). The between wild species and cultivars of Gladiolus (Ohri and
first Gladiolus hybrids were raised by Dean Herbert in Khoshoo, 1983b; Cantor and Tolety, 2011). Gladiolus
the 1820s, producing hybrids between various Cape palustris Gaudin and G. imbricatus L. indigenous to Europe

Volume 9, Number 2, May-August, 2020 121

 S.K. Datta

were recorded for Gladiolus species (Hamilton, 1980). As many wild species flower during winter and modern
Natural, interspecific hybridization between these species cultivars of G. x grandiflora flower in summer, storing
was possible due to their close genetic relatedness. the pollen is necessary to facilitate hybridization. In
Putative hybrids, G. palustris X G. imbricatus, has been addition, flower abortion of G. x grandiflora occurs in
reported in Poland (Kaminski, 2012; Cieslak et al., 2014). the winter because of low light intensity, and the species
Both species are cold-tolerant and their seeds and corms is not cold resistant (Imanishi, 1989). Winter-flowering
require a cooling period to germinate (Jogar and Moora, wild species are tolerant of low light intensity and cold
2008). Natural hybridization between G. palustris and G. temperatures. Such interspecific hybridization between
imbricatus was confirmed by chloroplast (psbA-trnH and G. x grandiflora and G. tristis was initiated to introduce
rpl32-trnL) DNA and nuclear ribosomal DNA (ITS1) some characters of the latter species, such as winter
sequences, AFLP markers and macro-, micro- flowering, cold tolerance, and fragrance into the former
morphological and reproductive characters (Szczepaniak hybrid. This cross combination has been reported in Israel
et al., 2016). CSIR-NBRI, Lucknow developed varieties with some miniature cultivars (Cohen and Barzilay, 1991).
(‘Manmohan’, ‘Manohar ’, ‘Mukta’, ‘Manhar’, This study demonstrated that long-term storage of pollen
‘Manisha’, ‘Mohini, ‘Urvashi’, ‘Neelima’, etc.) suitable of wild gladiolus is possible -20°C and that low
for cultivation in Indo-Gangetic Plains involving gladiolus temperature has a positive effect on interspecific crossing.
cultivar ‘Friendship’ (2n=60) and Gladiolus tristis (2n=30). These results suggest that interspecific hybridization
Breeding resulted in development of large number of between a modern cultivar and a wild species can be made
triploid (2n=45) hybrids (Sharga and Sharma,1984; more efficiently with these techniques and can be a useful
Sharma et al., 1988). In hybridization programme, seed tool to create new products for the cut flower market
setting is very important for intervarietal crosses. Different (Takatsu et al., 2002). Excellent parents can give poor
aspects like seed setting, early capsule maturity, highest progenies, since the phonotypical value of parents does
seed germination, higher number of seeds per capsule not permit a prediction of the value offspring. Experiment
and per cross were assessed in a number of cross was conducted to compare the morphological characters
combinations (Kumari et al.,2016). Interspecific of five genotypes and results indicated significant variation
hybridization between a modern cultivar of Gladiolus x amongst the genotypes with respect to morphological
grandiflora hort. (2n=60) and the wild species G. tristis characteristics as well as with yield, yield attributes and
L. (2n=30) was made to introduce characteristics of the plant height and white genotype has been mentioned as
wild species into the cultivated one. Gladiolus x the best planting materials (Hossain et al., 2011). Cantor
grandiflora is a summer-flowering species, and G. tristis and Chis (2009) developed two promising varieties
flowers in winter. The effect of storage temperature on ‘Candida Ali’ (early flowering cyclamen ‘Early Riser’ x
pollen viability was tested, as long-term storage of pollen early flowering pink ‘Priscilla’) and ‘Excelsa’ (pink
was necessary to facilitate crossing these two species. ‘Priscilla’ x move-cyclam ‘Speranta’). ‘Candida Ali’
Pollen of G. tristis could be stored at -20°C for  year, showed vigorous growth, early flowering, multiple middle
and was more practical than storage at -80°C. Air sized bright pink cyclamen florets with good simultaneous
temperature affected pollen tube growth, fertility, and fruit floweringand ‘Excelsa’ was very early flowering, red-
set in the cross between G. x grandiflora and G. tristis, coral floret. Gladiolus palustris is believed to be one of
and low temperatures (15°C–20°C) were best. The the rarest elements in the Polish flora and at the greatest
morphological data and flow cytometric analysis showed threat of extinction. AFLP studies of the genetic diversity
that the F1 plants were hybrids between G. x grandiflora of G. palustris from ex situ conservation at the Botanical
and G. tristis (Takatsu et al., 2002). Modern cultivars of Garden of Wroc³aw University, Poland revealed putative
G. x grandiflora were bred originally from only six species hybrids between G. palustris and G. imbricatus (Kamiñski,
(Barnard, 1972), and considerable genetic potential exists 2012; Cieœlak et al., 2014). Anthers of gladiolus open
for developing new kinds of gladiolus using wild species. and release their pollen a day or two before the stigma

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 Breeding of Ornamentals: Gladiolus

becomes receptive which is a constraint for inbreeding. improvement in ornamental and horticultural value of
Some admirable varieties have been created by inbreeding. gladiolus (Manzoor et al., 2018).Gladiolus is highly
Ervin H. Doerr developed ‘Bridesmaid’ (multi-coloured heterozygous which makes it an ideal material for
with intense ruffling) inbreeding ‘Boise Belle’ through 17 polyploidy induction that results in the production of new
successive generations (c.f. Paul H. Franklin, Fine forms in which one or few traits are improved without
Gardening). G. communis subsp. byzantinus appears to changing the whole genome.
hybridize with G. illyricus in southern Spain (Cantor and
Tolety, 2011). Spontaneous hybridization between G. Chromosome Studies
communis and G. italicus has occurred in Malta (Mifsud
Gladiolus is a representative ornamental where various
and Hamilton, 2013).
stages of transformation from wild to cultivated condition
largely affected by hybridisation and polyploidy are well
Mutation
preserved. A meaningful picture of origin and evolution
Appreciable amount of induced mutagenesis work has of garden gladiolus has been worked out on the basis of
been done on gladiolus using both physical and chemical cytogenetic mechanisms which is very helpful for further
mutagens. Mutation experiments generated huge amount improvement. Interspecific hybridization is possible
of basic and applied knowledge on different parameters regardless of ploidy or species level due to its flexible
of mutation and resulted in development of mutants with genome. Crosses between 2x and a 6x species produced
new flower colour, improved flower form, resistance to a highly heterozygous tetraploid hybrid. Many cycles of
lodging or disease, etc. Mutations in flower colour mostly further hybridisation involving some other species resulted
arise as chimera in gladiolus. The size of the mutant in amazing variability in present day cultivars (Ohri, 2013).
chimera varies from a narrow streak on a tepal to the Chromosomal studies on species and cultivars illustrate a
entire floret. Such chimeric mutant tissues cannot be range of ploidy from diploid (2n=30) to dodecaploid
isolated using the available propagation technique. Mutation (2n=180) but all the garden cultivars are tetraploid
in gladiolus can be established in pure form when mutated (2n=60). Meiotic behaviour and pollen stainability of
cells participate in corm/cormel formation in subsequent diploid and tetraploid taxa exhibited predominant bivalent
generations. Therefore, many new flower colour/shape formation along with pentavalent, quadrivalent, trivalent
mutants are lost. Management of such chimera and in and univalent. Studies revealed that the species native to
Cape Winter-Rainfall region (G. tristis, G. carneus, G.
vitro mutagenesis for induction of solid mutants have been
lapeirousioides, etc.) are diploid and those occurring in
well standardised in chrysanthemum. Direct shoot
Summer-Rainfall region and Tropical Africa (G.
regeneration will help to retrieve mutants from chimeric
oppositiflorus, G. primulinus, G. natalensis, etc.) range
tepals. So induced mutation combined with in vitro
from diploid (2n=30) to hexaploid (2n=90); while
technique is must in gladiolus to change one or a few
Mediterranean, European and West Asian species (G.
characters of an otherwise outstanding cultivar without
italicus, G. byzantinus, G. illyricus, G. segetum, G.
altering the remaining and often unique of the genotypes
communis, etc.) are all polyploidy, ranging from tetraploid
(Datta, 2014, 2017b).
to dodecaploid (Ohri and Khoshoo, 1985a,1985b; Cantor
Colchicine and Tolety, 2011; Valente et al., 2011). The majority of
the African species of Gladiolus are diploids (2n=2x=30)
Polyploidy (mixoploids + octoploids) and polyploidy plants whereas the European species are polyploids (2n=60–130;
were produced after treatment with different Cantor and Tolety, 2011; Valente et al., 2011; Delpierre
concentrations of colchicines (0.1%, 0.2% and 0.3%). and Du Plessis, 1973; Lewis et al.,1972; Ohri and
These putative polyploids may be helpful for further Khoshoo, 1985a; Tombolato et al., 2002).

Volume 9, Number 2, May-August, 2020 123

 S.K. Datta

Pollination Mechanism temperature stress of 8°C caused increased catalase (CAT)

activity and lower hydrogen peroxide (H2O2) levels in
For successful breeding, knowledge on pollen grain gladiolus, but such information has not resulted in breeding
fertility is an important pre-requisite character before for winter-hardy gladioli. Literature clearly indicates that
undertaking hybridization programme. Pollen fertility was majority of commercial gladioli have been developed for
tested for all germplasm collections (CSIR-NBRI, providing cut flowers in the summer. These varieties
Lucknow). It was interesting to note that 88% of cultivars
produce low-quality flowers when grown in subtropical
had pollen grain fertility above 50%. Maximum cultivars
and Mediterranean climates during winter (Magic and
had pollen fertility between 80% and 90% (Datta, 2001).
Cowperthwaite, 1954; McKay et al., 1981; Shillo and
Differential response to self-incompatibility, selfing and
Halevy, 1976a,1976b). Different environmental factors
crossing were studied on different species and garden
like low light intensities, short photoperiods, and the
cultivars. Experiments were conducted on pollination
combined effect of low irradiance with low night
studies in open pollination, natural cross-pollination,
temperatures, have been identified as the reasons for the
bagging, self-pollination and cross pollination and pod
failure to grow high-quality gladiolus during winter months
formation of each crossing for understanding the breeding
(Halevy et al., 1984; Shillo and Halevy, 1963, 1975,
system (Ohri and Khoshoo, 1981; 1983a, 1983b).
1976a,1976b, 1981). This motivated scientists to develop
Gladiolus is cross pollinated crop due to self-
cultivars to produce high quality flowers under winter
incompatibility. Long-term storage of pollen in gladiolus
conditions (Cohen and Barzilay, 1991).
is necessary to expedite artificial pollination for crossing
between spring-flowering species and summer-flowering
Disease Managements
species and varieties that bloom at different times. Pollen
of the gladiolus can be stored in plastic vials for longer Quite a large number of diseases cause serious damage
duration under freezer condition (-4oC) up to 11 weeks to gladiolus and floriculture trade suffers massive losses.
of storage without any alarming loss of viability (Johnson Extensive work has been carried out to ward off/minimise
et al., 2009; Kaur and Dhatt, 2019) and also in liquid the losses. About 20 pathogenic fungal species belonging
nitrogen (Balaram et al., 2000). As many wild gladiolus to 15 genera have been reported from all over the world
species flower during winter and modern cultivars of G.
on different varieties of gladiolus causing leaf spot to corm
x grandiflora flower in summer, pollen storage is necessary
rot. The details about the symptoms, main causal agents,
to facilitate interspecific hybridization. Takatsu et al.
extent of losses and the control measures of some of the
(2002) demonstrated that hybridisation between modern
major diseases (wilt, neck rot, corm rot, leaf and flower
cultivars and G. tristis was efficient at low air temperature
blight, viral diseases, Aster yellows; pest–thrips, cut worm,
(15°C–20°C) using pollen of G. tristis stored at -20°C.
leaf caterpillar, mite, mealy bug, etc.) under open field
This technique applicable to another wild species to breed
and greenhouse cultivation have been worked out and the
scented cultivars.
major approaches in controlling these diseases using
chemical, cultural methods and breeding for disease
Environmental Effects
resistance discussed (Datta, 2001; Sharma and
There are several environmental factors that affect the Bhattacharjee, 2002; Shanmugam et al., 2011; Nasir et
winter hardiness trait, including low temperatures, variable al., 2012). From gladiolus breeding stock few valuable
snow/ice cover, low light periods, and secondary invasion traits like resistance or immune to Botrytis gladiolorum,
by pathogens (Takatsu et al., 2002; Ali et al., 2016). Fusarium oxysporum f. gladioli, Curvularia trifolii, and
Winter hardiness is a desirable trait for gladiolus as well Stemphylium sp.; resistance to cold weather, flowering
as consumers for breeding. Gladiolus is a genus that has adaptation to short winter days, etc. were identified.
not been studied to any great extent in the area of winter Varieties (‘Florida pink’, ‘Picardy’, ‘Spic’ and ‘Span’)
hardiness. Bettaieb et al. (2007) found that low released during 1958 were very promising (Magie, 1960).

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 Breeding of Ornamentals: Gladiolus

Different diagnostic methods (based on RT-PCR, immuno- quality characteristics and helpful for colorant industries
diagnostics, molecular biology, DAS-ELISA, DTBIA, (Takemura et al., 2005; Islam, 2016). Twenty-three
ISEM, use of DNA/RNA probes, the polymerase chain anthocyanins have been detected from the flowers of
reaction, etc.) for detecting various viral diseases (Bean Gladiolus cultivars (Yatomi and Arisumi, 1968; Akaviaa
yellow mosaic virus (BYMV), Cucumber mosaic virus et al., 1981). 3-galactosides of kaempferol, quercetin,
(CMV), Arabis mosaic (ArMV), Broad bean wilt (BBWV), myricetin, laricitrin and syringetin as well as other
Tobacco mosaic (TMV), Tobacco necrosis (ToNV), flavonoids were found from the flowers of a wild species,
Tobacco black ring (TobBRV), Tobacco ringspot Gladiolus tristis. The major anthocyanins of the red
(TobRSV), Tomato ring spot (TRSV), Tomato spotted flowered cultivar ‘Shukuho’ and the pink ‘Naples’ were
wilt (TSWV), Soybean mosaic (SMV) and Strawberry different from the purple flower cultivars ‘Ariake’ and
latent ringspot, etc.) have been standardised and their ‘Blue Lagoon’. ‘Naples’ contained many other
management has been recommended (Raj et al., 2010; anthocyanins. White cv. ‘Saiun’ and yellow ‘Jacksonville
Katoch et al., 2004; Singh et al., 2015a, 2015b). Gold’ did not contain anthocyanin. The major anthocyanin
of the red ‘Shukuho’ was suggested to be a cyanidin
Mini Cultivar derivative. The major flavonoid was kaempferol. The
purple flower cultivars ‘Blue Lagoon’ and ‘Ariake’
Miniatures are extremely versatile because of their small
contained many flavonoids compared with other flower
plant size which can be grown easily in beds and most
colours suggesting that those flavonoids represent the
favourable in landscape plantings, lightweight, suitable for
more purplish colour as copigments (Scheffeldt and
density planting, reduced transport costs. A new group
Hrazdina, 1978). The six anthocyanidins in four types of
of miniature gladiolus was developed to meet the market
glycosylation were found in Gladiolus petals. Six
demand which was named ‘Orchidiola’ (orchid glads) by
monoglucosides appear in minute quantities, whereas any
the Israeli flower industry. Mini forms were developed
of the other 18 anthocyanins can serve as the major
from seedling selections from intercrossed of various
contributor to the coloration of Gladiolus petals (Akaviaa
ecotypes of the diploid species G. tristis and several
et al., 1981). Number of anthocyanins reported by
tetraploid commercial varieties (G. x grandiflorus).
different authors in G. x gandavensis varies as—six
Separate group of miniature varieties (‘Pixiola’) was
(Robinson and Robinson, 1931; Macek et al., 1946), two
developed in America (Barnard, 1972; Halevy et al., 1984;
on two hybrids (Robinson and Robinson, 1932,1934,
Cohen and Barzilay, 1991; Hort et al., 2012). The dwarf
Cohen et al., 1986), seven (Shibata and Nozaka, 1963),
gladioli, ‘Nanus’ hybrids’ (=G. nanus) is an interspecific
thirteen (Yatomi and Arisumi, 1968), sixteen (Arisumi and
hybrids from G. tristis, G. carneus, G. primulinus, and G.
Kobayashi, 1971). Seilleur (1977) studied quantitative
cardinalis (Goldblatt, 1996; Goldblatt and Manning,
relationship of the pigment between induced mutants of
1998).
the cultivar ‘Hawaii’ and its progenitor. Four types of
glycosylation of the six common anthocyanidins—
Pigment
pelargonidin, cyanidin, peonidin, delphinidin, petunidin and
Flower colour and its commercial value are determined malvidin—with the sugars glucose and rhamnose, were
by flower pigments, among others. Pigment composition reported in commercial tetraploid Gladiolus cultivars.
is very important as it provides adequate message for Pigments of each group may appear individually or in
practical breeding programme. The anthocyanin conjunction with pigments from either or both groups,
compositions in the flowers of different gladiolus all pigments contributing their share to the visible colour
genotypes were assessed and wide variation was observed in the petals. Flowers having a single pigment in their
about their total and individual anthocyanin constituents. petals were also found. The presence and absence, and
The results may be useful for various chemical breeding the relative amounts of specific pigments of each group
program improving desired organoleptic and nutritional affect the results of crosses between selected parents.

Volume 9, Number 2, May-August, 2020 125

 S.K. Datta

Specific parental clones containing a single pigment gladiolus in commerce, by out-crossing with ‘Mibloom’
produce offspring lines having no pigmental variation and and then recrossing those seedlings. Spencer selected more
little or no difference in the visible colour. This opens the than 150 lines with varying amount of fragrance (c.f. OE
possibility for F 1 hybrid production in gladiolus. Spencer, Zone10.com, Florida State Horticultural Society,
Understanding the inheritance of pigments determining 1960). One particular trait found in wild species is floral
colures is very important for application of chemical colour scent. Previously studied wild species and their major
analysis data for breeding. Gladiolus breeders should scent compounds are G. liliaceus (eugenol); G. alatus, G.
consider all these information for identifying parents when maculates, G. recurvus and G. tristis (linalool); G.
selectively breeding for colour. There is need to develop jonquilliodorus, G. orchidiflorus, G. patersoniae and G.
sound knowledge on different groups of pigments in scullyi (geraniol or geraniol acetate, nerol and citronellol);
existing varieties, their biosynthetic pathways, etc. and G. virescens (B-ionone; Goldbatt et al., 1998; Goldbatt
and Manning, 2002). Suzuki et al. (2008) analysed floral
Fragrant Gladiolus scent from 9 wild species and detected 20 scent
compounds of which 9 were benzenoids, 7 monoterpenes
Many original species that produced modern gladiolus and 4 sesquiterpenes. The species were divided into 4
were fragrant. These fragment species were crossed groups based on their major scent compounds like—
among themselves and with standard gladiolus. Crosses Lonalool/Benzenoid group (G. recurvus and G. tristis), Ione
produced varieties carrying qualities of modern gladiolus group (G. trichonemifolius, G. uysiae and G. watermeyeri)
and also the fragrance of the species. The maximum used and Ocimene/caryophyllene group (G. gracilis; Suzuki et
species was G. tristis. The main drawback to this natural al., 2008).
crosses is that the fragrant quality had been neglected
and lost during selection for flower size, colour and form. Molecular Characterisation
The size of floret and spike are small in the fragrant species
and also spring flowering which is the main bottle neck PCR-based techniques (RAPD, RFLP, SSR, STS, AFLP,
of breeding and producing normal Summer-flowering DAMD, ISSR, etc.) have been applied to study the genetic
gladiolus. But many common varieties still carry recessive variability, germplasm characterisation and relationships
strains of fragrance from their original ancestors. An early amongst gladiolus cultivars. The information generated
white with a red spot in the throat variety ‘Mibloom’ was will facilitate choosing the appropriate breeding program
to incorporate beneficial genes from genetically divergent
probably identified as a fragrant gladiolus. ’Incense’ a
cultivars in desirable genotypes lacking the particular trait
small pink with yellow throat variety was forerunner for
(Singh et al., 2015a, 2015b; Gonai et al., 2005; Pragya et
its fragrance. The next cultivar ‘New Era’ was identified
al., 2010a, 2010b; Dallavalle et al., 2002; Jingang et al.,
as slightly fragrant. ‘Frilled Fragrance’, a bud sport of
2008; Ranjan et al., 2010; Raycheva et al., 2011; Nasir et
‘New Era’ was introduced for sale. Another small ruffled
al., 2012).
rose pink variety ‘Summer Fragrance’ (a ‘Queen of
Bremen’ seedling) had light fragrance. ‘Mora’ (a ‘Shaylor’
Tissue Culture
seedling) is a light pink with a large contrasting red throat
that has an odour rather than a fragrance. All these fragrant Ziv et al. (1970) reported first micropropagation of
varieties were of unknown parentage or developed as gladiolus and subsequently the technique has been enriched
chance crosses. Under chance crosses category ‘Diadem’ for different species and varieties by a number of workers.
(‘New Era’ and ‘Vista Bonita’ parentage) was one of the Tissue culture procedure applied effectively in breeding
largest in the fragrant class. Another light fragrance variety programme to develop virus-free plants and for fast and
was ‘Gwen’ (a seedling of ‘Rose Marie Pfitzer’ and reliable micropropagation of elite clones. In vitro selection
‘Meanly’). A ruffled cream white with a rose dart in the at cellular level to assess genetic variation reduces time to
throat variety ‘Perfume’ is still one of the most fragrant breed and improve variety over conventional breeding

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 Breeding of Ornamentals: Gladiolus

method. Liquid culture and simple bioreactors systems et al., 2000). An Agrobacterium mediated transformation
were standardised to reduce production cost and to protocol using pre-wounded in vitro derived shoot tips
enhance multiplication rate. Efficient systems for the shoot bombarded with 1.6m naked gold particles by the biolistic
regeneration and rapid mass shoot propagation, shoot and delivery system has been developed (Babu and Chawla,
corm regeneration, ‘bioreactor’ system for production 2000). Molecular breeding is a good preference to develop
of uniform and vigorous corm-bearing shoots, high new flower colour and form using sense and anti-sense
frequency in vitro bud multiplication, cell suspension strategy or incorporation of new coloured genes in
cultures, selection of the gelling agent, standardisation of ornamentals. Molecular breeder should be very sensible
conditions for in vitro corm formation, etc. have been and choosy before starting such work considering the
optimised (Ruffoni et al., 2012; Choudhary et al., 2010; present diversity in floret colour in gladiolus.
Menon et al., 2016; Nhut et al., 2004; Sinha and Roy,
2002; Roy et al., 2006). DISCUSSION

Biotechnology The review is an important treatise that contains large

volume of information on different aspects of gladiolus.
New variety of bulbous ornamentals takes 5–10 years to It presents a complete scenario on multifarious aspects
be introduced into the market through natural vegetative of research on gladiolus, so that innovations made so far
propagation. Micropropagation is essential for can be used judiciously for the development of this sector.
development of large scale quality planting material and Creation of genetic variability is pre-requisite for
introduction of new variety within short period. To development of new variety and breeder should intelligently
overcome the restrictions of classical breeding application assess all up-to-date breeding techniques and knowledge
of biotechnology is a necessity for future ornamental for designing the breeding experiments. The breeder should
breeding. Molecular breeding is anticipated to have be conscious about the potential and the limitations of all
enormous potential for developing desired varieties. Cormel earlier experiments and reports and should consciously
derived cells were useful for inducing an embryogenic choose the knowledge which will provide accurate
cell suspension and in vitro selection of somaclonal guidelines for reaching the aims within reasonable time
variation (Remotti and Loffler, 1995). Regeneration of frame. Selection of desired characters began when wild
gladiolus has been standardised using gelling agent and species was brought under cultivation for domestication
kanamycin for successful plant transformation (Chauvin and improvement. The genetic evolution under
et al., 1999). Nhut et al. (2004) reported an economic in domestication is influenced altogether through different
vitro ‘bioreactor’ system for production of long term selection pressure and different objectives of man.
production of uniform and vigorous corm-bearing shoots. Hybridisation and introgression cycles have played very
Transgenic gladiolus plants have been developedvia important role in the diversification and evolution of
microprojectile bombardment of various regenerative Gladiolus L. (Barnard, 1972; Ohri and Khoshoo, 1983a,
tissues (sliced cormels, cell suspension and callus; Kamo 1983b; van Raamsdonk and de Vries, 1989; Goldblatt et
et al., 2005). Kamo (2003) reported highest levels of GUS al., 2001). It has been possible to produce a coherent
expression in callus, shoots, and roots of plants carrying picture of the various steps involved in the transformation
the bar-uid A fusion gene under control of the CaMV 35S of wild gladiolus to a cultivated one based on the available
promoter and in shoots and roots of greenhouses grown literature. Such a study has not only revealed the nature
plants that contained the rolD promoter. Transformation and extent of genetic evolutionary processes
protocol has been developed using embryogenic cells and accompanying domestication, but also help to chalk out
particle bombardment with gold particles coated with a steps for further improvement (Ohri and Khoshoo,
construct harbouring the gus reporter gene and the pat 1983a,1983b). It has been well proved that the genetic
(resistance to the herbicide Basta) selection gene (Loffler variability in different characters of any species is

Volume 9, Number 2, May-August, 2020 127

 S.K. Datta

influenced through breeding and recombination followed should not be restricted only for assessment of genetic
by selection. Such knowledge is very helpful in chalking diversity, identification of species specific markers and
out a breeding methodology in relation to the various traits phylogenetic relationship. Breeders must select and utilise
of the breeding system (Khoshoo, 1981). Review clearly closely and distantly related varieties to introgress new
indicates that we must change our mindset and go ahead and novel genes through selective breeding programme.
with selective breeding to develop specific desired As time proceeds, we have to change our routine research
phenotypic traits by choosing parents having desirable with a desire to make a fruitful and desirable difference
characters without conducting routine random breeding within short time. This state of positive mindfulness can
strategy. Such type of review is much needed in present be attained only through enhancing a scientific temper on
times in ornamentals (Datta,2017a, 2017b, 2018, 2019a, the basis of available work report, by which one can
2019b; Datta and Janakiram, 2015). It is particularly of evaluate what is right and desirable and what to do and
help to young scholars for whom access to original papers not do. As there is continuous need for modern breeding
of pioneering work on gladiolus is difficult if not programmes for novel characteristics and genes, gladiolus
impossible. Moreover, recent books also address this topic researchers should sensitise themselves regarding
rather superficially. preservation of wild relatives as well as those local
varieties.
CONCLUSION
ACKNOWLEDGEMENTS
Many publications on correlation studies are simple
I thank and acknowledge all professional colleagues/
repetition with routine findings and does not add much
scientists for their voluminous and interesting
new to the present knowledge. The information and
contributions in breeding of gladiolus. I sincerely thank
knowledge generated so far is a prelude for further
Dr. Manoj Nazir, Senior Scientist for his Facebook message
utilisation of promising genetically divergent cultivars of
on gladiolus. I acknowledge CSIR-National Botanical
gladiolus in the breeding programme to improve the
Research Institute, Lucknow where I did all my research
economically and industrially important traits of this one
on different ornamental crops on multidisciplinary aspects.
of the globally important floricultural crops. The present
available diversity in gladiolus is adequate for the future
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