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D. Prasath
ICAR-Indian Institute of Spices Research, Kozhikode, India
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Horticulture College and Research Institute, Tamil Nadu .Agricu/tural University, Coimbatnre - 647 003.
* Indian Institute of Spices Research, P. B. No. 7701, lI1arikunllu Post, Caticllt, Kerala - 673 012 .
Summary
High yielding varietiC's developed by plant brecdC'rs has resulted ill green revolution, caused self sufiicicncy in food production.
As the \vorld market develops mort-:> 10wards healthy cdting Jnd naturL'l1 products, the quality of the raw malerial from
agriculture beconws more criticaL The indiscriminatp use of synthetic colours for the food colouring has several harmful
effects. This necessitated our focus towards natural colours for food colouring, in general dnd development of high colour
eh i II i variet it'S for extraction of pL1prika oleore,,1 n, in partl cu la r. Capsicum breed Ing for qual ity has evolved through pure Ii I1C
.)election of local varietlcs to h,:brid devc)opment. Thl" review highlights achlpvements in breeding for high extractablE' colour
and pungency L1nd brccding strategies. The .)cope of the dlscu~.)ioll add resscs colour and pungency pri nci pies, genetic re50Uln~\
diverSity, specie::. compa1ibility, v,nlcty dnd hybrid development and biotechnological <)pplDaches.
I,ycollene
one of which contributes the fatty acid moiety Jnd the Capsaicinoid bios~nth(''Sis
Ranking ofvvorld CdpsicUlrJ bd ... ed 011 the pungc'Jlcy lc'v(,1 (Anon, 2003; AIlOIl, 200Scl; KUTllLll' ct al., 2006)
Iwai et a/. (1979) examined the fluctuation of pungent Pepper peroxidase isoenzyme B6, is capable of
principles of hot pepper fruits of Capsicum aonuwn oxidizing capsaicin and the phenolic precursors of
var. annuum cv. Karaystzubusta at different growth capsaicin, with caffeic acid and ferulic acid being the
stages after flowering. Capsaicinoid was detected 20 b,'st substrates among them and the basic peroxidase
days afterflowering, which reached maximal level at isoenzyme B6 is also located in cell walls (Bernal et
around 40 days after flowering and then decreased ai, 1993). Bernel et al. (1995) suggested that this
gradually. Capsaicin was always the major isoenzyme may be involved in the insolublisation of
component, occupying approximately 60 per cent of phenylprop,moids precursors. These results support
the capsaicinoids with a capsaicin: dihydrocapsaicin the biochemical evidence which poillts to the existence
ratio of 1.36:1.71. of an oxidative competitive sink for phenylpropanoid
intermediates of capsaicin biosynthesis, which
Enzymes involved in the capsaicin metabolism
probably competes with cap sal< in itself in the cell wall
Phenylalanine ammonia lyase (PAl) is the enzyme that of Capsicum anlll/l/m.
catalyses the first step in the pathway ,,,ing L
Genetic diversity
phenylalanie as the substrate and rendering cinnamic
acid, which is subsequently converted into r Consideration and assessment of genetic variability
coumaric, caffeic, and ferulic acids by the action of existing in a population is the basic reql/irement for
cinnamic acid - 4 - hydroxylase (CA 4 H), r-coumaric (lny effective crop improvempnt programlllE'.
acid - 3 - hydroxylase (CA,H), and caffeic acid -0 Introgression of different desir<lble traits spread ill the
methyl transferase (CAOMT) respectively. The enzymes diverse genotypes into single genotype is desirable for
that participate in the transformation of ferulic acid any crop. The diversity for pungency ,md colour value
into vanillin and vanillylamine are unknown. Finally, of the Indian chillies has been studied by different
the enzymatic condensation of vanillylamine and 8 research workers and are summarized in the following
methyl - 6 - nonenoic acid is carried out by tables.
'capsaicinoid synthetase (CS)' (FuJiwake c( ,1/., 1980).
Genetic resources and conservation
Fujiwake et al. (1982) determined that the biosynthetic
The International Pldnt Genetic Resources Institute
site of capsaicinoids is in the placenta of the fruit and
(IPGRI, 1983) has mad" dforts to organize pl'lIlt
produced by the cinn,'mic acid pathway. Iwai ct a/.
explorations and both working and base collections
(1979) suggested that capsaicinoids production
of Capsicum genetic resources. In 1986, the Asian
incredse with maturity until a maximum and then
Vegetable Research and Development Center
decreased by rapid turnover and degradation of upto
(,....VRDC) was invited to mainta'" a b,lCkup of global
60 per cent. In general, the fruit tissue showed a low
Capsicum collections (Poulos, 1994). In India, til"
activity of PAL, C\H and CS during the period from
systematic collection, charMterization and
o to 22 days alter flowering and a peak of activity for conservation of Capsicum germplasm for qUdl ity were
all the enzymes at the time of maximum growth in
undertaken by both by Indian Council of Agricultural
length of the fruit (Alejo and Peralta., 1993)
Research (Indian Vegetable Research Institute,
FUJiwake et a/. (1982) observed that the capsaicinoids Varandsi; Indi'ln Institute of Spices Research, Cdlicl/t;
progressively accumulated during the development of Indian Agriculture Research Institute, Regional Station,
the fruit, reached maximum of 120 mg/g in field grown Katrain and Inelidn Institute of Horticulture Research,
after 45-50 days after fruit set dnd then started to Bangalore) and Sate Agricultural Universities
decrease gradually. The caps,licinoids started to (University of Agricultural Research, Dharwad ,'nd
accumulate just before the increase in length ceased Tamil Nadu Agricultural University, Coimbatore).
(between day 20 and 35) and active accumulation took
A prospect for the future is to enhance germ plasm pool
place OVl'r a 15 day period (day 25 to 40) with an
development with other domesticated species of
average increase of 333 mg day" per fruit. After day
Capsicum, particularly frolll Me'ico, Central and
40, the level of capsaicinoids per fruit remained
South America and parts of Asia and Africa wher"
virtually constant (Sukrasno and Yeoman, 1993).
species in addition to C. annUl/rrJ are cultivated.
EXTRACTABLE COLOUR AND PUNGENCY IN CAPSICUM 5
Heterosis for pungency and colour value coordinate regulation (Curry ei al., 19991 QTL interval
analysis for individual and total capsaiclnoid COlltent
Although Capsicum annuum tolerate inbreeding,
identified a major QTL, termed cap, which explained
various workers have dernonstrated considerable
34 - 38 per cent of the phenotypic variation for this
extent of heterosis for pungency and colour val ue.
trait in two growing environments (Blum ei al. 2(03).
Deshpande (1933) recorded less pungency in hybrids
The molecular linkage map of C locus has been
than that of their parents. Absence of heterosis for thiS
prepared and a pungency ,,'Iated gene has been found
pungency was ob,erved by Sharrna and Saini (1977).
to be located on chromosome 2. Based on sequences
Later, pOSitive heterosis were reported by (Sekar, 1984;
of caspsaicinoid synth,lSe (CS), sequence
Anandanayaki 1997; Tanki, 1999; Hernavathy, 2000;
characterized amplified region (SCAR) markers have
Do,hi and Shukla, 2000 and Sathiyarnurthy, 2002),
been developed and their usefulness in mrly detection
averaging between 8.84 and 98.08 per cent. Prasath
of pungent or non-pungent genotypes have been ,1[SO
(2005) reported that the hPlerosis for extr<lCtable colour,
demonstrated (Lee E't al., 2005).
which ranged frorn -47.84 to 32.11 per cent over
better p,lIent. The variety, KDC - \, a hybrid derivative The absence 01 capsanthin and capsorubin in yellow
of cross between By,ldagi and C. frutescens was fruits correlates with the lack of expres"on of CCS
released from LIAS, Dharwad, Karnataka. It bears semi (capsan\hin-capsorubin synthase) enzyme in yellow
wrinkled red fruits with high colour and low pungency. truits (Bouvier pi a/., 1994; Houln" et a/., 1994). Co
dominant DNA markers for the identification of red
Gene action
and yellow-fruited genotypes at seedling stage hdve
It has long been known that single dorninant gene, C, been developed (Popovsky and Paran, 2000).
controls the presence or absence of pungency in the Metabolic engineering of ClpsiculJl, with r"gard to
fruits (Blum et al., 20021. However, in the pungent enhanced cap::'dicin or carotenoid pdthway gpne has
types, the degree of pungency is quantitatively been dernonslTdted in several systerns, inclucJing rice
inherited and highly influenced by environment (Ye ei al., 20(0). However, there is a need to get ,\
(Zewdie and Bosland, 2000). Studi"s conducted so high pigment and low pungency product from
far have indicated that maturPd red colour is domin,'nt Capsicum, which will be of valup for
over yellow colour and is controlled by a single gene pharamaceuticLlls, consumer purposps and as a food
(Y) and later it was reported that colour compounds co'lourant. Attempts to get both a placenta-sill'ciflc
(carotenoids) is under the control of four different genes expression of caps,\icinoids and a fruit wall Si,,'cific
(y, c" c l and c') with ppistatic interaction (Hurtado expression of carotc'noids is also needed. Such
Hernandez and Smith,1985); Shifriss and Pilov,ky , developmellts need to be completed for dised'"
1992). Popov sky and Paran (2000) reported mature resistance, which would result in till' overall
fruit co!our is under influence of three independpnt improvement of the ClpsiculTI lor pre and post h,lIvest
pair of genes (eI, c2 and Y).Th,' presence of dominant applications for augmt'nting qualiLJtive and
alleles at these three loci results in red mature frUits, quantitative output of the products from Capsicum.
while recessive al[eles give white mature fruits.
Conclusion
Biotechnological approaches
Achievements in Capsicum breeding havl' culmilldted
The growth of molecular biology and instrumentation in the release of specialized rnarket types alld the
paved th" way for the I,ltest breeding approach which advent of hybrid varieties. Ldnd races and facultative
is easy and quick. This also proved to be very important self poll inatpd vMieties also conti nue to oce upy mdlor
approach for quality brpeding at the present and future production areas in developing countries. The recent
conditions. Genes coding for enzymes in capsaicinoid discovery of new medicinal properties of carotenoids
biosynthesis pathway have been isolated (Curry ei al. and capsaicinoids of CapsicUl77 could be of gredt
1999; Kim <'t a/. 2001 and Aluru ei al 20(3). A study potential to become evermore versatile crop in world
of placental transcript [evels for the suite of structural agriculture, Thus, an interndtional focus is requirpd
genes thought to be involved in capsaicin biosynthesis on the development of Capsicum for high colour ,'nd
demonstrated a generdll correlation between transcript low or nil pungency through germplasm, genetics and
level and the degree of pungency, consistent with breeding. As far is India is concerned, majority of the
EXTRACTABLE COLOUR AND PUNGENCY IN CAPSICUM 7
chilli area IS caniined to tropics and thereiore, our capsaicin content of Capsicum fruits at different stages
attention should be the development oi paprika type oi maturity. Lloydia. 31: 272-274
chilli genotypes suitable to tropical conditions. Also, Ben Chaim, A., Paran, I., Grube, R., Jahn, M., Wijk R.and
systematic attempt has to be made in germplasm Pelema" I. (2001). QTL mapping oiiruit related traits
collection and evaluation oi low pungent, high colour in pepper (Capsicum anlluum). Theor. Appl. C('llet.
types such as Bydagi and tomato chilli. Furthermore, 102: 1016-1028.
identiiication oi hybrid combinations with high Bennett, D. J. and Kirby, G. W. (1968). Constitution and
economic yield coupled with high colourdnt and biosynthesis oi c<,("aiclll. /. Chem. Soc. 442-446.
oleoresin recovery would help in increasing the Bernal, M. A., Calderon, A. A., Pedreno, M. A, Munoz, R.
colourant and oleoresin yield per unit area substantially. R., Barcelo ,~. and Caceres, F. M. (1993). 1hc
subcellular localization of iso-peroxidases in Cap~icum
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EXTRACTABLE COLOUR AND PUNGENCY IN CAPSICUM 9