Biotechnology of eggplant - Université...

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3 Biotechnology of eggplant

4 V. Kashyapa, S. Vinod Kumara, C. Collonnierb, F. Fusaric,5 R. Haicourb, G.L. Rotinoc, D. Sihachakrb, M.V. Rajama,*

6 aPlant Genetic Manipulation Group, Department of Genetics, University of Delhi

7 (South Campus), Benito Juarez Road, New Delhi 110021, India

8 bBat 360, Morphogenese Vegetale Expeimentale, Universite Paris Sud, 91405 Orsay Cedex, France

9 cIstituto Sperimentale per L ‘Orticoltura. Via Paullese 28, 26836 Montanasao Lombardo-Milano, Italy

10 Received 1 December 2000; received in revised form 20 June 2002; accepted 24 July 2002

11

12 Abstract

13

14 Eggplant (Solanum melongena L.) is an important vegetable crop grown in various tropical and

15 temperate parts of the world. There is a wide genetic diversity in the cultivated as well as the wild

16 species of eggplant. Cultivated varieties of eggplant are susceptible to a wide array of pests and

17 pathogens as well as to various abiotic stress conditions. In contrast, the majority of wild species are

18 resistant to nearly all known pests and pathogens of eggplant and thereby are a source of desirable

19 traits for crop improvement. Tissue culture protocols for organogenesis, somatic embryogenesis,

20 anther culture and protoplast culture have been well established for the eggplant. Somatic hybridisa-

21 tion has also been attempted for transferring useful genes from wild species to the cultivated plants

22 through protoplast fusion. However, the information on genetic engineering and molecular biology of

23 eggplant is very limited. Transgenic eggplants for insect resistance, for the production of parthe-

24 nocarpic fruits and abiotic stress tolerance have been accomplished. However, transgenics of

25 eggplant are yet to be developed for improvement of other agronomic traits, including disease

26 and pest resistance, and quality and shelf life of fruits. Molecular markers to complement traditional

27 breeding programs are being developed for genome mapping of agronomic traits. The present review

28 summarises efforts to improve eggplant genetics with an emphasis on the use of biotechnology to

29 introgress genes from wild species into cultivated eggplant.

30 # 2002 Published by Elsevier Science B.V.31

32 Keywords: Solanum melongena; Plant regeneration; Anther culture; Review; Protoplast fusion; Transgenic

33 plants; Aubergine34

Scientia Horticulturae 1846 (2002) 1–25

* Corresponding author. Tel.: þ91-11-4679866; fax: þ91-11-6872437.

E-mail address: [email protected] (M.V. Rajam).

1 0304-4238/02/$ – see front matter # 2002 Published by Elsevier Science B.V.

2 PII: S 0 3 0 4 - 4 2 3 8 ( 0 2 ) 0 0 1 4 0 - 1

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35 1. Introduction

36 Solanum melongena L. ð2n ¼ 24Þ, commonly known as eggplant, aubergine, guinea

37 squash or brinjal, is an economically important vegetable crop of tropical and temperate

38 parts of the world (Table 1). It is a good source of vitamins and minerals (particularly iron)

39 making its total nutritional value comparable with tomato (Kalloo, 1993). Eggplant has

40 been used in traditional medicines (Khan, 1979). For example, tissue extracts have been

41 used for treatment of asthma, bronchitis, cholera and dysuria; fruits and leaves are

42 beneficial in lowering blood cholesterol.

43 Eggplant is a native of the Indian sub-continent, with India as probable centre of origin

44 (Gleddie et al., 1986a). The medicinal and economic value of eggplant is found in Sanskrit

45 literature (Kalloo, 1993; Hinata, 1986; Khan, 1979). Indo-Burma, China and Japan are the

46 secondary centres of eggplant origin (Gleddie et al., 1986a). Eggplant has been cultivated

47 in Asia for over 1500 years. Arabs introduced eggplant to the west during the 15th century

48 (Hinata, 1986). Eggplant germplasm resources and collections have been well documen-

49 ted, evaluated and conserved throughout the world (Sarathbabu et al., 1999).

50 Eggplant has been divided into three main types egg-shaped (S. melongena var.

51 esculentum); long slender shaped (S. melongena var. serpentium) and dwarf type (S.

52 melongena var. depressum) (Kalloo, 1993). There are also many wild species of eggplant

53 that carry many economically important genes. Wild eggplant species show resistance to

54 important pests that affect commercial eggplant production (Table 2).

55 Eggplant improvement using conventional approaches have been directed towards

56 improvement of agronomic traits such as fruit size, weight and shape, and resistance to

57 diseases and pests. Conventional breeding in eggplant has been reviewed (Kalloo, 1993).

58 Therefore, main focus in the present review will emphasise information on the biotech-

59 nological aspects of eggplant improvement with only, brief information included on

60 breeding aspects. Thus, the review covers the past, present and future prospects of eggplant

61 improvement, which is being undertaken worldwide.

62 2. Eggplant cytology

63 Cytological studies on eggplant species have been instrumental in classification of the

64 plant. Although, the basic chromosomal number ðn ¼ 12Þ is same in all the varieties and

Table 1

World wide production of eggplanta

Area harvested (ha) Average yield (kg/ha) Production (mt)

World 1,313,903 169,514 22,272,454

Asia 1,234,002 167,272 20,641,440

Africa 45,260 178,451 807,668

Europe 27,487 242,695 242,695

North and Central America 6,589 227,015 149,580

South America 425 137,647 5,850

Oceania 140 58,571 820

a FAO, 2001 report.

2 V. Kashyap et al. / Scientia Horticulturae 1846 (2002) 1–25

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O65 species, the chiasma frequency during diplotene and diakinesis show varied bivalents

66 (Kalloo, 1993). The cytology and chromosome number of many diploids, autotetraploids,

67 autotriploids, amphiploids and androgenic haploids have been well studied in eggplant

68 (Kirti et al., 1984; Rao and Baksh, 1979; Siliak-Yakovlev and Isouard, 1983). The

69 karyotypic studies by Venora et al. (1992) have shown that haploidy has no effect on

70 the shape and size of the chromosomes. The karyotype comparisons between wild S.

71 sisymbriifolium Lam., S. torvum Sw. and cultivated S. melongena show that S. sisym-

72 briifolium Lam. is phylogenetically distant to S. torvum Sw. and S. melongena, which are

73 closer to each other than the former (Russo et al., 1992).

74 The cytological studies carried out earlier are now well supported by the evidence

75 obtained through DNA fingerprinting. Sakata et al. (1991) initiated phylogenetic studies in

76 eggplant by comparing isozymes and pattern of cpDNA between wild and cultivated

77 eggplants. They comprehend that scarlet eggplants should be treated as single species (S.

78 aethiopicum L.) derived from single wild species (S. anguivi Lam.). In a similar study,

Table 2

Sources of wild eggplant species resistant to pathogens/pests

Pathogens/pests Resistant wild species References

Fungal wilts

Fusarium wilt S. indicum, S. integrifolium,

S. incanum

Yamakawa and Mochizuki (1979)

Verticillium wilt S. caripense, S. periscum,

S. scabrum, S. sisymbriifolium,

S. torvum

Sakata et al. (1989), Fassuliotis and

Dukes (1972), Petrov et al. (1989)

Bacterial wilts S. integrifolium, S. torvum Yamakawa (1982), Sheela

et al. (1984)

Fruit rot

Phomopsis vexans S. gilo, S. integrifolium Ahmad (1987)

Cercospora solani S. macrocarpon Madalageri et al. (1988)

Fruit and shoot borers

L. orbonalis S. xanthocarpum, S. khasianum,

S. integriifolium,

S. sisymbriifolium

Chelliah and Srinivasan (1983),

Sharma et al. (1980), Khan

et al. (1978)

Root-knot nematodes

Meloidogyne spp. S. sisymbriifolium, S. torvum,

S. aethiopicum, S. warscewiczii

Fassuliotis and Dukes (1972), Ahuja

et al. (1987), Di Vito et al. (1992),

Daunay and Dalmasso (1985),

Hebert (1985)

Spider mite S. macrocarpon, S. integriifolium,

S. mamosum, S. pseudocapsium,

S. sisymbriifolium

Schalk et al. (1975), Schaff

et al. (1982)

Other pests/pathogens

Aphis gossypii S. sisymbriifolium, S. mammosum Sambandam and Chelliah (1983)

Tetranychuns urticae S. macrocarpon Schaff et al. (1982)

Egg plant mosaic

virus (Tymovirus)

S. hispidum Rao (1980)

Mycoplasma S. hispidum, S. integrifolium Rao (1980), Khan et al. (1978)

V. Kashyap et al. / Scientia Horticulturae 1846 (2002) 1–25 3

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79 Sakata and Lester (1994) showed that S. melongena and wild species S. incanum L. are very

80 close in phylogeny than the wild species S. marginatum L. fil. (Fig. 1). Later, the studies by

81 allozyme patterns and RAPDs also show that cultivated S. melongena, the weedy form S.

82 insanum L. and the wild species S. incanum L. are phylogenetically very close (Karihaloo

83 and Gottlieb, 1995; Karihaloo et al., 1995). Isshiki et al. (1994, 1998, 2000) conducted

84 segregation studies for seven isozyme loci of five enzymes in cultivated and wild species of

85 eggplant. They have suggested three linked pairs for the isozymes studied and thus these

86 seven loci can be used as genetic markers. Even seed storage proteins have also been used

87 to characterise the wild and cultivated eggplant species (Menella et al., 1999).

88 3. Interspecific hybridisation using embryo rescue in eggplant

89 The quantitative and qualitative breeding efforts in eggplant have resulted in introduc-

90 tion of many improved varieties into cultivation. Resistance to pests and pathogens has

91 been well identified in many wild species, which has been summarised in Table 2. Most of

92 the wild species carrying useful traits belong to the sub-genus Leptostemomocus of family

93 Solanaceae (Daunay and Lester, 1988; Daunay et al., 1991). Interspecific hybrids between

94 wild and cultivated species have been successful in only a few cases (McCammon and

95 Honma, 1983; Sharma et al., 1980, 1984; Daunay and Lester, 1988). Table 3 summarises

96 interspecific hybridisation from conventional crosses.

Fig. 1. Dendrogram given by cluster analysis of cpDNA. Weighed pair-group method of hierarchical and

unmatching distance was employed for clustering. u: S.0050 and S.1780. v: S.2026, S.2029, S.2369. S.1382,

S.1781, S.1398, S.2055, S.0931, S.1793, S.1512 and S.1750; w: S.1554, S.0657, S.1961 and DMO; x: S.1397 (S.

lichtensteinii); y: S.1518, S.1692 and RNL337 (S. lichtensteinii); z: S.0256 and S.0396 (Adapted from Sakata

and Lester, 1994. Chloroplast DNA diversity in eggplant (S. melongena) and its related species S. incanum and

S. marginatum. Euphytica 80: 1–4).


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Table 3

The interspecific crosses between the cultivated S. melongena and various wild species

Parents for crosses Status of hybrids References

S. melongena � S: indicum F4 plants obtained Rao and Kumar (1980), Rao and

Rao (1984)

Partially fertile plants Krishnappa and Chennaveeraiah

(1965), Rajasekaran (1968),

Narasimha Rao (1968), Rangaswamy

and Kadambavanasundaram

(1973a,b, 1974a,b)

S. melongena � S: sodomeum Fertile F1 plants Tudor and Tomescu (1995)

S. melongena � S: macrocarpon F1 and F2 fertile plants Schaff et al. (1982)

Sterile plants Rajasekaran (1961), Wanjari (1976),

Gowda et al. (1990)

S. melongena � S: khasianum Successful F1 hybrids after

embryo rescue

Sharma et al. (1980, 1984)

S. melongena � S: aethiopicum F1 hybrids obtained after

embryo rescue

Ano et al. (1991)

Fertile hybrids Ignatova (1971)

S. melongena � S: insanum F1 hybrids obtained after

embryo rescue

Ali and Fujieda (1990)

Fertile hybrids Swaminathan (1949), Mittal (1950),

Babu Rao (1965)

S. melongena � S: gilo F1 hybrids obtained Ali and Fujieda (1990)

Sterile F1 hybrids Nasrallah and Hopp (1963),

Omidiji (1981)

Partially fertile hybrids Narasimha Rao (1979)

S. melongena � S: hispdum Sterile hybrids Rao (1980)

S. melongena � S: torvum F1 hybrids after embryo

rescue, but the plants had very

low fertility (1.73%)

McCammon and Honma (1983),

Blestsos et al. (1998)

Unsuccessful cross Rao and Rao (1984)

S. melongena � S: integrifolium F1 hybrids Rao and Baksh (1979)

Sterile F1 plants Berry (1953), Fukumotoh (1962),

Rao and Baksh (1981), Kirti and

Rao (1982), Khan et al. (1978)

Partially fertile hybrids Hagiwara and Iida (1938, 1939),

Tatebe (1941), Miwa et al. (1958),

Kataezin (1965), Narasimha

Rao (1968), Ludilov (1974)

S. melongena � S: sisymbriifolium Embryo rescued hybrid

plantlets did not survive

Sharma et al. (1984)

Sterile hybrids obtained

after embryo rescue

Blestsos et al. (1998)

S. melongena � S: xanthocarpum F1 hybrids sterile Rajasekaran (1968, 1971), Sarvayya

(1936), Hiremath (1952)

Partially fertile hybrids Swaminathan (1949)

S. melongena � S: grandiflorum Partially fertile hybrids Ramirez (1959)

S. melongena � S: cumingii Fertile plants Capinpin et al. (1963), Fukusawa

(1964)

S. melongena � S: zuccagnianum Sterile F1 hybrids Rajasekaran and

Sivasubramanian (1971)

S. melongena � S: surattense Sterile F1 hybrids Rao and Rao (1984)

S. melongena � S: indicum Sterile F1 hybrids Rao and Rao (1984)

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97 In several cases, crosses were only successful if in vitro embryo rescue was employed.

98 Sharma et al. (1980) was successful in obtaining fertile hybrids between S. melongena and

99 S. khasianum Clarke. through embryo rescue, but only when the wild eggplant S.

100 khasianum Clarke. was selected as the female parent. The successful hybrids were obtained

101 when naked embryos (25 days old) were cultured on Nitsch and Nitsch (1969) medium.

102 However, in the present case no hybrids could be obtained with ovule culture. They also

103 achieved partial success in obtaining hybrid plants between S. melongena and S. sisym-

104 briifolium Lam., when the latter was taken as the male parent and the excised torpedo

105 staged embryos were cultured on Nitsch and Nitsch (1969) medium. However, in the latter

106 case the plants could not survive for very long time and died (Sharma et al., 1984). Daunay

107 et al. (1998) have been successful in developing 22 hybrids between S. melongena and wild

108 species of eggplant. In another study, Blestsos et al. (1998) through embryo rescue

109 developed hybrids with S. torvum and S. sisymbriifolium Lam. The hybrids were obtained

110 by culturing immature ovules on MS medium for 50 days. Thereafter, the embryos

111 dissected from these ovules were cultured on MS medium in dark for 10 days and then

112 transferred to light. They have further characterised the hybrids morphologically and by

113 isozyme studies. The F1 hybrids obtained after embryo rescue developed fruits only when

114 used as a female parent in backcrosses. The successful interspecific crosses have been

115 obtained with only few wild species. In such attempts, the hybrids have been developed

116 through embryo rescue. However, such hybrids have either been sterile or have had very

117 low pollen fertility (Sharma et al., 1980, 1984; McCammon and Honma, 1983; Blestsos

118 et al., 1998). This may be due to pre- and post-pollination effects (Kalloo, 1993; Narasimha

119 Rao, 1979; Omidiji, 1979; Sharma et al., 1980, 1984).

120 4. Organogenesis

121 Organogenesis has been successfully achieved in cultivated and wild varieties as well as

122 their hybrids (Fig. 2; Table 4). Fassuliotis (1975) was first to report regeneration in S.

123 sisymbriifolium Lam., a wild species of eggplant. Stem parenchyma cells were isolated and

124 cultured on Linsmeier–Skoog (LS) medium supplemented with 6-(g,g-dimethylallyla-

125 mino)-purine (2iP) and indole-3-acetic acid (IAA). Other wild species in which regenera-

126 tion studies have been carried out include S. xanthocarpum Schrad. & Wendl. (Rao and

127 Narayanaswami, 1968), S. aviculare Forst. f., S. gilo Raddi. (Gleddie et al., 1985a;

128 Kashyap et al., 1999), S. khasianum Clarke. (Bhatt et al., 1979; Kowalozyk et al.,

129 1983), S. indicum auct. non L. p. pte. and S. torvum Sw. (Gleddie et al., 1985a; Kashyap

130 et al., 1999).

131 Kamat and Rao (1978) reported shoot regeneration from hypocotyl segments of S.

132 melongena and the F1 hybrids in presence of cytokinins, kinetin and zeatin. The study by

133 Fassuliotis et al. (1981) on cell suspensions culture raised from pith callus in eggplant have

134 shown appearance of green nodules in presence of IAA and 2iP which fail to develop

135 further. Such cultures when transferred to medium containing ascorbic acid or antiauxin p-

136 chlorophenoxy isobutyric (PCIB) acid developed into organised shoots. The study by

137 Allicho et al. (1982) has also shown that presence of cytokinin is necessary for shoot

138 differentiation in eggplant. Adventitious shoots have been formed by BAP, zeatin or kinetin


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139 supplemented media (Gleddie et al., 1983; Mukherjee et al., 1991; Sharma and Rajam,

140 1995a). Sharma and Rajam (1995a) have shown that hypocotyl explants yield higher

141 adventitious shoots than cotyledons or leaves. They have shown that morphogenetic

142 response also varied within single explant and follows a basipetal pattern where the apex

143 region is better responding than the basal region. Recently, use of thidiazuron (TDZ) has

144 shown to enhance shoot organogenesis; the leaves and the cotyledons respond best to TDZ

145 (Magioli et al., 1998). Scoccianti et al. (2000) has recently reported the correlation between

146 the organogenesis from the cotyledon explants of eggplant and hormone modulated

147 enhancement of biosynthesis and conjugation of polyamines (PAs).

148 5. Somatic embryogenesis

149 Somatic embryogenesis (SE) in eggplant has been reported from stems, hypocotyl,

150 leaves, cell suspension, isolated protoplasts and roots (Matsuoka and Hinata, 1979; Gleddie

151 et al., 1983; Fobert and Webb, 1988; Kalloo, 1993; Sharma and Rajam, 1995a,b; Yadav,

152 1997; Yadav and Rajam, 1997, 1998). The first report of SE in eggplant was by Yamada

153 et al. (1967) in MS medium supplemented with IAA, whereas Matsuoka and Hinata (1979)

154 obtained somatic embryos from hypocotyl explants on MS medium supplemented with a-

155 naphthaleneacetic acid (NAA) (8 mg/l).

156 The studies on SE carried out so far reveal that NAA enhances somatic embryo

157 differentiation. However, its optimal concentration varies with the initial explant. Leaf

158 explants required 2–6 mg/l of NAA for SE differentiation while hypocotyls required 6–

159 10 mg/l (Matsuoka and Hinata, 1979; Gleddie et al., 1983; Sharma and Rajam, 1995a).

Fig. 2. Organogenesis in a wild species of eggplant (Solanum indicum). Induction of shoot regeneration (A),

excised individual shoot on rooting medium (B) and rooted in vitro shoot (C).


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Table 4

Organogenesis in various species of eggplant

Species Explant Basal

medium

Growth regulators (mM)

Callus formation Shoot formation Root Formation References

S. melongena Hypocotyl MS 0.6–11.4I AA þ 4.4 BA 0.6–11.4 IAA þ 4.4 BA 0.6–11.4 IAA þ 4.4 BA Kamat and Rao (1978)

0.6–11.4 IAA þ 2.3–4.7 Kn 0.6–11.4 IAA þ 2.3–4.7 Kn 0.6–11.4 IAA þ 4.7–9.3 Kn

2.8–11.4 IAA þ 2.3 Zn 2.8–11.4 IAA þ 2.3 Zn 2.8–11.4 IAA þ 2.3 Zn

5.4 NAA þ 4.4 BA

5.4 NAA þ 4.7 Kn

4.5 2,4-D þ 4 BA

S. melongena Hypocotyl MS 4.3–86 NAA 4.3–86 NAA 4.3–86 NAA Matsuoka and Hinata (1979)

4.3–86 BA 4.3–86 BA 4.3–86 BA

S. melongena Cell suspension

from pith

MS 11 IAA þ 4.5 2,4-D þ4.5 Kn

0.57–28.5 IAA þ37–74 2iP

49–74 2iP þ 0.01–1.0

Ascorbic acid

Fassuliotis et al. (1981)

S. melongena Hypocotyl MS 1.8–9 2,4-D 1.8–9 2,4-D Allicho et al. (1982)

Cotyledons MS 1.8–9 2,4-D 1.8–9 2,4-D

Leaf MS 1.8–9 2,4-D 1.8–9 2,4-D

S. melongena Leaf explants MS 0.05–0.4 TDZ Magioli et al. (1991)

S. melongena Hypocotyl MS 11.1 BA þ 2.9 IAA 1/2 MS Sharma and Rajam (1995a)

Cotyledons MS 11.1 BA þ 2.9 IAA 1/2 MS

Leaf MS 11.1 BA þ 2.9 IAA 1/2 MS

S. torvum Leaf MS 22.62 2,4-D 0.46–23.23 Kn 3.91 2,4,5-T Gleddie et al. (1985a)

0.39–19.57 2,4,5-T 0.46–22.81 Zn 28.54 IAA

0.54–26.85 NAA 22.19 BA

0.49–24.73 NOA 13.99 BARI

4.92–24.61 2iP

S. torvum Leaf protoplasts Modified KM 11.4 IAA þ 9.1 Zn Guri et al. (1987)

S. Khasianum Leaf discs MS 10 NAA þ 1 & 10 BA 0.1 NAA þ 10 BA 10 NAA þ 1 BA Kowalozyk et al. (1983)

10 NAA þ 1 IAA þ 10 BA 0.1 NAA þ 1 IAA þ 10 BA 10 IAA þ 1 Zn

10 IAA þ 10 Zn 10 IAA þ 10 BA 10 IAA þ 0.1 Zn

1 IAA þ 10 Zn 1 IAA þ 10 Zn

0.1 NAA þ 1 IAA þ 10 Zn 0.1 NAA þ 1 IAA þ 10 Zn

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10 IAA þ 1 & 0.1 Zn

S. Khasianum Leaf MS 4.5 2,4-D 4.44 BA 0.45 2,4-D Gleddie et al. (1985a)

0.39 2,4,5-T

0.54–4.37 NAA

5.7–28.5 IAA

0.53–5.29 IPA

S. sisymbriifolium Stem parenchyma

cells

LS 29.82–119.28 2iP þ0.28–85.62 IAA

Fassuliotis (1975)

S. sisymbriifolium Stem explants MS 0.45–22.62 2,4-D 4.56–45.2 Zn 4.52 2, 4-D Gleddie et al. (1985a)

19.57 2,4,5-T 4.44–22.19 BA 3.9 2,4,5-T

0.54–26.85 NAA 4.92 2iP 0.54–26.85 NAA

5.7–28.54 IAA

S. gilo Leaf MS 4.52 2,4-D 22.81 Zn Gleddie et al. (1985a)

5.37–26.85 NAA

4.95–24.73 NOA

S. aculeatissimum Leaf MS 0.46–23.23 Kn 0.45–22.62 2,4-D Gleddie et al. (1985a)

0.46–22.81 Zn 0.54–5.36 NAA

0.44–22.19 BA 0.49–4.94 NOA

0.1–2.8 BARI 0.57–5.71 IAA

S. aviculare Leaf MS 0.45–22.62 2,4-D 23.23 Kn 5.71–57.08 IAA Gleddie et al. (1985a)

0.39–19.57 2,4,5-T 0.46–22.81 Zn 5.29 1PA

5.37–26.85 NAA 0.44–22.19 BA

4.95–24.73 NOA 0.28–13.99 BARI

5.71–28.54 IAA 4.92–24.61 2iP

S. aviculare Shoot MS 4.52 2,4-D 0.46–23.23 Kn Gleddie et al. (1985a)

0.39–19.57 2,4,5-T 0.46–22.81 Zn

5.37–26.85 NAA 0.44–4.44 BA

28.54 IAA 2.98–14.91 2ip

26.43 IPA

S. xanthocarpum Shoot explants White 27.14 2,4-D Rao and

Narayanaswami (1968)

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160 Synthetic seeds have also been developed by encapsulating somatic embryos with sodium

161 alginate and calcium chloride (Lakshmana Rao and Singh, 1991; Mariani, 1992).

162 Somatic embryogenic response in eggplant was also genotype-dependent. Sharma and

163 Rajam (1995a) compared genotype, explant type and position on SE using various

164 Indian cultivars. Leaves and cotyledons showed higher SE induction compared to

165 hypocotyls. Interestingly, significant differences for morphogenetic potential were

166 noticed within a single explant (hypocotyl); and the terminal hypocotyl segmentex-

167 hibited better embryogenetic potential than the median segments (Sharma and Rajam,

168 1995a,b). Saito and Nishimura (1994) obtained non-vitrified somatic embryos from cell

169 suspension cultures.

170 Momiyama et al. (1995) investigated the expression of mRNA products during early

171 stages SE through differential display studies. Eight new mRNA products were identified

172 from 4-day-old tissue cultures. However, the expression of only one of the products

173 remains uniform throughout early stages (initial 10 days) of SE. Further, they have cloned

174 the cDNA corresponding to this expression product. The product cloned has been found to

175 share high similarity with Arabidopsis cDNA clone. This clone of Arabidopsis is a

176 homologue to ausprressor of Wilm’s tumor.

177 Hitomi et al. (1998) showed that the auxin type used for SE could impart somaclonal

178 variation. Using morphological features of somatic embryogenic raised plants they showed

179 that the somaclonal variation was higher in plantlets obtained with hormone NAA

180 compared to 2,4-D. This is accounted from the variations in the plant habit (tall or dwarf)

181 and leaf shape (narrow or thick leaves). The plants obtained with hormone NAA had all

182 types of variations, i.e. plants with larger leaves (4.5%), dwarf habit (4.5%), narrow leaves

183 (3.5%), larger leaves and dwarf plants (6.6%) and narrow leaves with dwarf habit (1.4%),

184 whereas the plants regenerated through 2,4-D had only one variation, i.e. they were having

185 narrow leaves with dwarf habit (11.8%).

186 PAs are important regulators for SE in eggplant. During SE in S. melongena cotyledon

187 explants treated with NAA exhibited high free putrescine (Put) and spermidine (Spd) in

188 comparison to those without NAA (Fobert and Webb, 1988). The temporal changes in

189 cellular PA concentrations were closely associated SE process (Sharma and Rajam

190 1995a,b; Yadav and Rajam, 1998). Embryogenic potential of explants collected from

191 different regions of leaf (apical and basal discs) was related to the spatial distribution of

192 endogenous PA levels (Yadav and Rajam, 1997). It has been shown that the apical region of

193 leaf explants of eggplant having a high SE potential have higher PAs than the basal region

194 that yields lesser somatic embryos (Sharma and Rajam, 1995b). Further, Put promoted

195 somatic embryo formation by sixfold. However, Spd and Spermine (Spm) have no

196 stimulatory effect. When the Put inhibitor difluoromethylornithine (DFMO) was added

197 externally, there was a reduction in Put and Spd titres and a corresponding reduction in SE.

198 The inhibitory effects of PA inhibitor difluoromethylarginine (DFMA) on SE could be

199 restored by adding exogenous Put (Yadav and Rajam, 1997). Yadav and Rajam (1998)

200 studied the temporal regulation of SE by modulating PA biosynthesis in eggplant. Put

201 levels increased during early stages of SE and the judicious time and dosage of PA/PA

202 biosynthesis inhibitor (DFMA) allowed the modulation of PA metabolism and further

203 regulation of SE. Such studies may be helpful in promotion and induction of plant

204 regeneration via SE in morphogenetically poor and recalcitrant species, respectively.


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205 Indeed such an approach has been used successfully in rice for promotion of regeneration

206 from poorly responding genotypes (Shoeb et al., 2001). The above studies clearly suggest

207 that PAs have an intricate regulatory role in SE.

208 6. Anther culture

209 Studies with anther culture have mostly been conducted for cultivated eggplant (S.

210 melongena) with goal of obtaining double haploid parents for conventional breeding

211 (Rotino, 1996). The double haploid plants have been successfully used in conventional

212 breeding programmes to obtain pure lines faster than selfed inbreds. Double haploid plants

213 are homozygous at all loci, and this may help to study the genetic basis of quantitative traits

214 by overcoming the problems associated with the environmental variations. The double

215 haploid parents have proven to be useful for breeding plants with useful agronomic traits,

216 such as high yield, disease resistance, earliness, abiotic stress tolerance and other

217 characters in numerous crops (Jacobsen and Sopory, 1978; Uhrig and Salamani, 1987;

218 Waari, 1996).

219 Raina and Iyer (1973) were first to report plant regeneration from anther culture in

220 eggplant. They regenerated homozygous diploid (double haploids) plants through callus

221 developed from anthers cultured at uninucleate pollen stage that were previously treated

222 with colchicine. Haploid plantlets were also obtained from the Research group of Haploid

223 breeding (1978) and Isouard et al. (1979) a year later. Dumas De Vaulx and Chambonnet

224 (1982) did an extensive work to improve the development of androgenic haploids. They

225 showed that high temperature ð35 � 2 �CÞ incubation of anthers under dark conditions for

226 the first 7–8 days improved the efficiency of haploid plant formation. A combination of

227 both auxin and cytokinin was essential during early stages of anther culture. Similarly,

228 Rotino et al. (1987) showed that haploid plant regeneration was affected by genotype,

229 temperature, culture conditions, hormones and anther stage. A higher temperature governs

230 the shift of the microspores from gametophytic stage to sporophytic stage. The regenera-

231 tion medium supplemented with same cytokinin as in the induction medium would

232 enhance the anther response (Rotino, 1996).

233 The effect of somaclonal variations on agronomic traits of embryogenic and androgenic

234 colchicine-treated double haploid lines was also investigated (Rotino et al., 1991). Plant

235 height, fruit shape and yield were some of the features affected by somaclonal variations.

236 Miyoshi (1996) cultured isolated microspores on Nitsch Lichter Nitsch (Litcher, 1982)

237 medium supplemented with NAA (0.5 mg/l) and BA (0.5 mg/l) initially for 4 weeks. The

238 calli obtained were transferred to MS medium supplemented with zeatin (4 mg/l) and IAA

239 (0.2 mg/l) for plant regeneration. The efficiency of haploid regeneration was additionally

240 improved by preculturing anthes or micorspores to induce callus formation prior to

241 inducing planlet regeneration (Gu, 1979; Miyoshi, 1996).

242 Anther culture was successfully employed to reduce the ploidy of eggplant somatic

243 hybrids between S. melongena, and wild species (S. integrifolium and S. aethiopicum gr.

244 Gilo) (Rotino et al., 2001; Rizza et al., 2001, 2002) to the diploid status and the

245 corresponding plants may be more easily crossed with the diploid cultivated eggplant

246 for transferring useful traits.


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247 7. Protoplast culture and somatic hybridisation

248 Plant regeneration from protoplasts has been achieved in both cultivated and wild

249 species of eggplant. The protoplast culture and somatic hybridisation would be useful in

250 overcoming the pre- and post-fertilisation breeding barriers encountered during conven-

251 tional breeding. Further, protoplast cultures are excellent means for understanding

252 cytological and ultrastructural changes during cell growth and differentiation, behavioural

253 patterns of plastids and mitochondria (Fournier et al., 1995). For cultured eggplant (S.

254 melongena), protoplasts isolated from mesophyll cells grew best using both cytokinin and

255 auxin (Sihachakr and Ducreux, 1987). However, protoplasts isolated from petioles and

256 stems showed better regenerating potential compared to cells isolated from lamina

257 (Sihachakr and Ducreux, 1987). In addition to hormone, the regeneration of plantlets

258 from protoplasts was highly genotype specific (Gleddie et al., 1983). An alternate pathway

259 for regeneration from protoplasts is via SE (Gleddie et al., 1983).

260 In wild species regeneration response is varied between species. Gleddie et al. (1985a)

261 have compared plantlet regeneration between S. sisymbriifolium Lam., S. torvum Sw., S.

262 gilo Raddi., S. aviculare Forst. f., S. aculeatissimum Jacq. and S. khasianum Clarke. They

263 found the best regeneration in S. aviculare Forst. f. and S. sisymbriifolium Lam., a minimal

264 response in S. gilo Raddi. and no regeneration in other species tested. However, Guri et al.

265 (1987) were successful in establishing plantlets regenerated from protoplasts from S.

266 torvum Sw. The protoplasts and protoplast obtained calli of S. torvum Sw. regenerated

267 better on modified Kao Michayluk (KM) medium supplemented with cytokinins and auxin

268 in comparison to MS medium as reported by Gleddie et al. (1985a) earlier.

269 Somatic hybridisation using protoplast fusion has facilitated transfer of useful traits in

270 eggplant between species that are sexually incompatible (Table 5). One of the main

271 advantages of somatic hybridisation over sexual hybridisation is that hybrids with novel

272 cytoplasmic and nuclear traits can be developed by hybridising distantly related species of

273 economic importance.

274 The first successful somatic hybrid was developed between cultivars of S. melongena

275 and S. sisymbriifolium Lam. through polyethylene glycol (PEG)-mediated protoplast

276 fusion (Gleddie et al., 1986b). The hybrid flowers were light purple in colour and displayed

277 floral abnormalities like separate petals, which was absent in either of the parents. Since

278 then somatic hybrids have been developed by fusing protoplasts of cultivated varieties with

279 S. aethiopicum L. group aculeatum (Daunay et al., 1993), S. torvum Sw. (Guri and Sink,

280 1988a) and S. khasianum Clarke. (Sihachakr et al., 1988). Interspecific somatic hybrids

281 were also developed by fusing protoplasts of S. melongena with S. nigrum L. to transfer

282 herbicide (atrazine) resistant trait from S. nigrum L. is indicated by Southern analysis (Guri

283 and Sink, 1988b). The intergeneric asymmetric somatic hybrids have been developed

284 between interspecific tomato hybrid (Lycopersicon esculentum Mill. and L. pennellii

285 (Corr.) D’Arcy [EP stands for is the hybrid between Lycopersicon species esculentum (E)

286 and pennellii (P)]) and eggplant (Liu et al., 1995). The somatic hybrids were obtained

287 through organogenesis showed flower abscission. In another similar study Samoylov and

288 Sink (1996) have shown that the regeneration ability of asymmetric somatic hybrids

289 depend upon the irradiation dose. The asymmetric hybrids were developed by exposing the

290 chromosomes of donor genome to different doses of gamma rays (100, 250, 500, 750 and


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O291 1000 Gy). It was found the hybrids exposed to 100 Gy gamma rays was a tetraploid (4n

292 ploidy), whereas exposure to other doses formed hybrids between 5n and 9n ploidy. The

293 radiation dosage determined the ploidy level and the DNA content of these hybrids.

294 Protoplast fusion has been achieved through electrofusion or PEG. The protocols for

295 both have been standardised (Sihachakr et al., 1993). The basal medium for protoplast

296 culture and regeneration medium has been Kao and Michayluk (KM) medium supple-

297 mented with hormones, namely, 2,4-D, NAA, BAP or zeatin in combination or alone (Kao

Table 5

Production of somatic hybrids from protoplast fusion in eggplant

Parents selected Culture response Reference

S. melongena (Black

BeautyÞ � S: sisymbriifolium

26 hybrid lines were obtained

mostly aneuploids having

chromosome number close to 48

Gleddie et al.

(1985b, 1986b)

S. melongena

ðDourgaÞ � S: khasianum

83 somatic hybrids obtained.

Most hybrids were tetraploids (48)

and few were aneuploids

Sihachakr et al. (1988)

S. melongena

(Black BeautyÞ � S: torvum

10 somatic hybrids were well

established and chromosome

number ranged from 46 to 48

Guri and Sink (1988a)

S. melongena (Black

BeautyÞ � S: nigrum

Only two somatic hybrids

could be obtained

Guri and Sink (1988b)

S. melongena

ðDourgaÞ � S: torvum

19 somatic hybrids were obtained

having chromosome number

ranged from 46 to 48

Sihachakr et al. (1989a)

S. melongena

ðDourgaÞ � S: nigrum

Only single somatic hybrid

could be obtained which was

aneuploid with 2n close to 96

Sihachakr et al. (1989b)

S. melongena

ðShironasuÞ � N: tabacum

Somatic hybrids were not

successful since only green shoots

could be obtained from two

somatic hybrid colonies

Toki et al. (1990)

S. melongena � S: macrocarpon The developed hybrids were

highly sterile and there was

failure to set seed

Gowda et al. (1990)

S. melongena � Lycopersicon spp. Only two hybrids with leaf like

primordia were obtained

Guri and Sink (1991)

S. melongena

ðDourgaÞ � S: aethiopicum

35 somatic hybrids were obtained

of which 32 were tetraploids, one

was hexaploid and two mixoploids

Daunay et al. (1993)

S. melongena (Black

BeautyÞ � hybrid of L. esculentum

and L. pennellii (EP)

These were asymmetric hybrids.

Only four hybrids could

be obtained

Liu et al. (1995),

Samoylov and Sink (1996)

S. melongena � S: torvum They selected 12 somatic hybrids

to be incorporated in the

breeding programs

Jarl et al. (1999)

S. melongena � S: aethiopicum 30 hybrid plants were obtained.

All hybrids were fertile. Of these

16 were tested tolerant against

bacterial wilt

Collonnier et al. (2001)


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298 and Michayluk, 1975; Sihachakr et al., 1993). The regenerated somatic hybrids were

299 obtained through callus.

300 The somatic hybrids have been well characterised using isozyme patterns, morphology

301 and cultural behaviour, molecular and physiological studies to confirm hybridization

302 (Gleddie et al., 1986a,b; Daunay et al., 1993; Sihachakr et al., 1993; Guri and Sink, 1988b;

303 Sakata et al., 1991; Filippone et al., 1992). Flow cytometry has been used to establish

304 ploidy level (Sihachakr et al., 1993; Filippone et al., 1992).

305 Many useful traits from wild species like resistance to fungal and bacterial wilts, and

306 nematodes have been maintained in the somatic hybrids. Recently, somatic hybrids

307 between S. melongena, S. aethiopicum L. group aculeatum and gilo have been developed

308 and found to be highly resistant to bacterial wilt caused by Ralstonia solanacearum

309 (Collonnier et al., 2001).

310 8. Genetic engineering

311 Successful eggplant genetic transformation was achieved as early as 1988 (Guri and

312 Sink, 1988a,b) using Agrobacterium-mediated genetic transformation with the cointegrate

313 vector- pMON 200 harbouring nptII gene. This was followed by a series of successful

314 attempts to improve the transformation protocol using gene constructs with nptII as a

315 selection marker and several reporter genes like gus (b-glucouronidase), cat (chloram-

316 phenicol acetyl transferase) and luciferase (Table 6). Though, a number of reports describe

317 transformation protocols, a comprehensive study to optimise factors that dramatically

318 affect transformation (explant type, genotype and cultural conditions) is lacking. A recent

319 study has made some efforts to standardise transformation protocol taking into account the

320 influence of antibiotics and growth regulators (Billings et al., 1997). More recently, an

321 efficient transformation protocol was developed and used for the generation of transgenic

322 eggplants tolerant to abiotic stresses by the introduction of bacterial mtlD gene (Prabhavati

323 et al., in press).

324 The insecticidal protein gene Bt(CryIIIb) from Bacillus thuringensis was used to

325 develop transgenic insect resistance in eggplant (Rotino et al., 1992b). Plants developed

326 were resistant to Colorado Potato Beetle (Arpaia et al., 1997). Field trials showed that

327 resistant transgenic lines had significantly higher fruit yield (Arpaia et al., 1998; Acciarri

328 et al., 2000). There was no significant difference in colonies of non-target insects like green

329 peach aphid (Myzua persicae Sulz.), flea beetle (Altica sp.), poatao tuber moth (Phthor-

330 imacea operculella) and lacewings (Chrysoperla carnes Stephens) in transgenic and

331 control fields (Acciarri et al., 2000). Introduction of a synthetic Bt(CryIAb) gene to

332 eggplant provided protection against fruit borer—Leucinodes orbonalis (Kumar et al.,

333 1998).

334 Parthenocarpic eggplant transgenics have been successfully developed by transferring

335 iaaM gene from Pseudomonas syringae pv. savatanoi driven by the DefH9 promoter from

336 Antirrhinum majus. Transgenic plants developed fruits from both pollinated and unpolli-

337 nated flowers in comparison to the untransformed control plants, which formed fruits only

338 when pollinated. In case of transgenic plants, seedless fruits were obtained only from

339 unpollinated flowers (Rotino et al., 1997). In a greenhouses trial transgenic parthenocarpic


UNCORRECTED PROO

Table 6

Development of eggplant transgenics through genetic engineering

Marker gene Reporter gene Useful gene Accomplishment Reference

nptII and hptII Successful Agrobacterium-mediated genetic transformation Guri and Sink (1988a,b)

nptII luciferase Agrobacterium-mediated transformation using callus cultures Komari (1989)

nptII Transgenic plants obtained from cotyledonary and leaf explant Filippone and Lurquin (1989)

nptII cat Development of efficient protocol for transformation using binary vector Rotino and Gleddie (1990)

nptII gus – Transformation in wild species of S. integrifolium Rotino et al. (1992a)

nptII gus – To study segregation of nptII gene in progenies of transgenic plants Sunseri and Rotino (1992)

nptII gus Efficient protocol for transformation through SE Fari et al. (1995b)

nptII gus Bt(CryIIIB) The introduced gene had low expression in transformed plants Chen et al. (1995)

nptII Transformed in S. gilo, a wild species of eggplant Blay and Oakes (1996)

nptII – Bt(CryIIIB) Introduction of modified Bt-gene into eggplant Colorado Potato

Beetle (CPB). The transgenics were highly resistant to CPB

Arpaia et al. (1997)

nptII gus Bt(CryIIIB) The increase in transformation efficiency with phytohormones

(TBZ and 2iP) and antibiotics (Kanamycin and Augmentin)

Billings et al. (1997)

nptII Modified Bt(CryIIIA) Introduction of modified Bt-gene in eggplant Hamilton et al. (1997)

nptII iaaM–DefH9 Introduction of gene responsible for parthenocarpy. The

seedless fruits developed as result of introduction of this gene

Rotino et al. (1997)

nptII Modified Bt(CryIII) The modified gene confirmed resistance in wild species

S. integrifolium and cultivated eggplant

Iannacone et al. (1997)

nptII Synthetic Bt(CryIAb) Introduction of synthetic Bt-gene. The expression of insecticidal

protein gave significant protection against fruit borer (L. orbonalis)

Kumar et al. (1998)

nptII uidA Synthetic Bt(CryIIIA) 69% of transgenic plants were resistant to neonate larvae and adult

CPB. The segregating F1 population also showed resistance the CPB

Jelenkovic et al. (1998)

nptII Bt(CryIIIB) Study of impact of developed transgenics on environment.

The resistant lines were safe for all non-target organisms

including mammals and environmentally safe

Arpaia et al. (1998),

Acciarri et al. (2000)

nptII DefH9–iaaM Winter cultivation in unheated greenhouse resulted in significant

increment in winter production, improvement of fruit quality

and reduction of cost

Donzella et al. (2000)

nptII mtlD Introduction of mannitol-1-phosphodehydrogenase conferred

tolerance to osmotic stresses induced by salinity and drought

Prabhavati et al. (in press)

V.

Ka

shya

pet

al./S

cientia

Ho

rticultu

rae

18

46

(20

02

)1

–2

51

5

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340 plants were more productive both natural parthenocarpic hybrid and induced partheno-

341 carpic plants using phytohormone sprays (Donzella et al., 2000). More recently eggplant

342 transgenics tolerant to abiotic stresses like salinity, drought and chilling have been

343 achieved by introduction of bacterial mannitol-1-phospho dehydrogenase (mtlD) (Prab-

344 havati et al., in press).

345 9. Molecular markers in eggplant

346 Plant breeding programmes all over the world are developing molecular markers through

347 RFLP, RAPD and AFLP (Tanksley et al., 1989; Williams et al., 1990; Vos et al., 1995).

348 The RAPD, RFLP and AFLP studies have also revealed distinction between wild and

349 cultivated plants of eggplant. These have very effectively shown genetic diversity and

350 relatedness among various eggplants (Isshiki et al., 1998; Mace et al., 1999; Kashyap et al.,

351 1999, 2000; Kashyap, 2002). Recently Nunome et al. (1998) have constructed linkage map

352 of eggplant. Their aim is to identify molecular markers linked to important agronomic traits

353 especially those resistant to bacterial wilt caused by R. solanacearum.

354 An eggplant RFLP genetic linkage map for qualitative and quantitative traits has been

355 constructed from an F2 segregating population derived from an interspecific cross with S.

356 linnaeanum Hepper & Jaeger (Frary et al., 2000). More recently, markers linked to

357 eggplant fruit shape and colour was identified in a molecular linkage map based on RAPD

358 and AFLP (Nunome et al., 2001). Therefore, the application of molecular assisted selection

359 (MAS) may also be applied in breeding of eggplant. The markers can function as probes for

360 the disease resistant traits. Further this may facilitate the identification and isolation of the

361 genes of interest faster. The genome mapping with probes and markers can help in studying

362 both Mendelian and non-Mendelian inheritance of genes. However, such studies in

363 eggplant are in their infancy.

364 10. Conclusions and future prospects

365 Biotechnology has opened up new vistas for crop improvement. Biotechnological tools

366 like in vitro propagation, genetic engineering and molecular biology has helped over-

367 coming constraints of conventional breeding, and identification and introduction of useful

368 genes that confer resistance to pests and diseases, and tolerance to abiotic stresses in

369 eggplant. Although some developments in the field of biotechnological applications has

370 taken place, the full potential is yet to be exploited for improvement of eggplant.

371 Plant regeneration studies have contributed much to the crop, as there are extensive

372 studies on regeneration via organogenesis and SE. The fundamental aspects of plant

373 regeneration have been addressed by many studies and considerable information has been

374 gathered at the cellular and molecular level. The understanding of specific metabolic

375 pathways directly or indirectly involved in plant morphogenesis has helped in under-

376 standing and improving the regeneration potential of eggplant genotypes. Improvement of

377 organogenesis and SE by manipulating the PA metabolism has opened new vistas not only

378 in eggplant tissue culture but also tissue culture of other crops. Further, the studies on


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379 differential display of transcripts during SE hold better hope in understanding the

380 molecular mechanisms that control the process of SE. The protoplast culture and somatic

381 hybridisation has been extensively studied in eggplant with the prime aim to introgress

382 valuable agronomic traits from wild species to cultivated eggplant. However, the sterility

383 among the somatic hybrids obtained through conventional breeding and other genetic and

384 physical barriers still remains a hurdle. Further efforts are needed to overcome this and to

385 utilise the potential of this method.

386 The marker studies like RAPD, RFLP and AFLP are not been exploited fully for eggplant

387 breeding. These studies need to be strengthened for developing phylogenetic maps and

388 molecular markers for disease and pest resistance as well as for other agronomic traits.

389 Plant transformation procedures have been well established and are being utilised for

390 producing eggplant transgenics. However, genetic engineering has not yet been utilised to

391 its full potential for eggplant improvement as the existing studies have been confined

392 mainly to the development of insect resistant transgenics and transgenics for partheno-

393 carpic fruit development as well as tolerance to abiotic stresses. Thus, such studies need to

394 be further exploited so as to address the development of transgenic plants tolerant to the

395 wide array of environmental stresses, and fungal and bacterial pathogens. Post-harvest

396 quality improvement, increased shelf life, nutritional quality improvement, increased

397 water use efficiency and delayed senescence are a few of the many areas which have

398 to be addressed by genetic engineering techniques. The prospects of eggplant improvement

399 appear brighter with the advent of biotechnology tools. The areas that need to be

400 strengthened in eggplant research are genetic engineering and molecular biology. There

401 is already an international concern and many collaborative projects have been launched

402 which include European Commission collaborative projects on the development of

403 valuable breeding materials of eggplant resistant to bacterial and fungal wilts as well

404 as root nematodes diseases through somatic hybridization (Grant No. ERBIC

405 18CT9701878), the Indo-French collaborative research programmes for the development

406 of transgenic eggplants resistant to fungal and bacterial diseases by the over-expression of

407 pathogenesis-related protein genes (Grant No. 1803-1), the INDO-USAID Plant Germ-

408 plasm Project, and European Union research programme for the molecular analysis of the

409 cultivated eggplant and wild relatives.

410 Uncited references

411 Anonymous, 2000; Guri and Sink, 1998c; Kashyap and Rajam, 1999.

412 Acknowledgements

413 The research work on eggplant biotechnology has been generously supported by the

414 European Commission (Grant No. ERBIC 18CT9701878), the Indo-French (Grant No.

415 1803-1) and the Department of Biotechnology, New Delhi (Grant No. BT/R&D/08/40/96).

416 Senior Fellowships from the Council of Scientific and Industrial Research, New Delhi to

417 VK and SVK is gratefully acknowledged.


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