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FIG
(
Fic us caric a
)
AkathSingh, JaiPrakash,P.R.MeghwalandS.A.Ranpise
1. INTRODUCTION
Fig (Ficus carica) is one of the oldest
known fruit trees in the world. Turkey
produces 26% of the world’s figs and
Egypt,Iran,Greece,Algeria,andMorocco
together produces around 70% of the
world’s fig production (FAO, 2006). In
India,itscultivationismostlyconfinedto
westernpartsofMaharashtra(Pune)and
Gujarat, Uttar Pradesh (Lucknow and
Saharanp ur), K arna t a ka (Bellary,
CnitradurgaandSrirangapatna)andTamil
Nadu(Coimbatore).Figisespeciallywell
adapted toMediterranean climates, with
cool winters and hotdry summers, but
can be grown in more humid regions
including thetr opics and subtropics. Fig
is ex tremely dr ought toler ant once
established and commercial production is preferr ed in dry hot summer but needs regula r
irrigation for achieving higher yields.
Figs can be eaten fresh, dried or canned and are often used in preparation of jam.
As a fresh fruit, it has a luscious taste. Fruits have been prized over centuries for the
medicinalanddietaryproperties.Figisahighlynutritiousfruit.Itisrichincalories(269),
proteins,andcalcium(higher thanmilk), ironandhighest fibrecontent. Fighasnutritive
index of 11,as against 9, 8 and 6 for apple, raisina nd date, respectively. The chemical
composition andflavour of figvaries with the cultivar. Thetotal sugar contentof fresh
CONTENTS
1. Introduction...............................................149
2. Domestication ............................................150
3. Taxonomy ................................................... 150
4. CentersofOrigin/CentersofDiversity...... 150
5. ObjectiveofImprovement .........................151
6. Cytogenetics............................................... 151
7. InheritanceofPattern ...............................151
8. ProblemsinBreeding ................................ 152
9. FloralBiology ............................................. 152
10. DifferentSpecies .......................................158
11. TypesandCultivars ................................... 158
12. CropImprovementMethods......................161
13. VarietalWealth .........................................166
14. FutureThrust ............................................. 174
12
150Breeding of Un derutilized F ruit Crops
figis16 %andofdriedis52%.Theedibleportionoffreshfigs(100g)containsmoisture
(88.1%), protein (1.3g), fat (0.2%), minerals (0.6%), fibre(2.2%), carbohydrate (7.6%),
calcium (35mg), phosphorus (22mg), iron (0.6mg), vitamin A (80 IU), vita min C2 mg,
calorific value80andthiamine(0.1mg). Theedibleportionofdriedfigs(100g)contains
moisture(23.0%),protein(4.3g),fat(1.3%),minerals(2.4%),fibre(5.6%),carbohydrate
(69%),calcium(200mg),phosphorus(77mg),iron(4mg),vitaminA(100IU),vitamin
C2mg,calorificvalue306 andthiamine(0.1mg).Figisavery poorsourceofVitamin
C but very rich in sugar next to da tes. The latex from fig contains (ficin) rennin, which
has3to100timesmilkclottingcapacityascomparedtoanimalrenninpreparedformcalf
stomach mucosa. Fig leaves are used medicinally for their diuretic, demulcent, emollient
and antihelmintic properties. Three new constituents viz., calotropenyl acetate, lupeol
acetate and oleanolic acid were isolated by Ahmed et al (1990). Fig is valued for its
laxative properties and is used in the treatment of skin infection. The fruits help to
maintainacidalkalibalance ofthebody.Latexisusefultocoagulate milk(Morslietal.,
1985).Manymedicinalvirtueshavebeenascribedtothefig.Itisconsidereda restorative
foodwhichhelpsinquickrecoveryafterprolongedillness.Itremovesphysicalandmental
exertion and endows the body with renewed vigor and strength. It is an excellent tonic
for the weak people who suffer cracks in lips tongue and mouth.
2. DOMESTICATION
ItwasperhapsfirstbroughtintocultivationinthesouthernpartsoftheArabianPeninsula
byatleast3000BC.ItlaterspreadintoIran,SyriaandTurkeyandintoalltheMediterranean
countries.DuringtheageofexplorationfollowingthediscoveryofAmericabyColumbus,
the fig was taken to most subtropical areas of the western hemisphere. Dushevskii and
Kazas (1985) described two forms of Ficus carica from the vicinityof Mangup Kale,
a settled area on Mt. BabaDag inCrimea abandoned in 1783. They are thought to be
descendants of figs cultivated on the plateau from the 12th century.
3. TAXONOMY
It belongs to family moraceae, genus Ficus, is a monoecious, deciduous tree or large
shrub. The subgenus Eusyce to which Ficus carica belongs is characterized by having
unisexual flowers only and gynodioecism (Storey, 1975). The inflorescence is unique,
consisting of a syconium (the fig), which encloses many unisexual flowers that can be
accessedviatheostiolebypollinatingwaspsthat maygivethetruefruits.Tinypedicellate
drupletsusuallycalledthe‘seed’(Storey,1975).Morphologically Ficuscarica isconsidered
gynodioeciousbutfunctionallydioecious.
4. CENTERS OF ORIGIN /CENTERS OF DIVERSITY
It is thought to be a native to southern parts of Arabian Peninsula, Italy, the Balkan
Peninsula and the former USSR (Tutin, 1964). Accor ding to ElRay and Llacer (1995),
Fig(Ficus carica)151
the fig originated in the east Mediterranean region (Turkey, Syria, SaudiArabia), from
where its cultivation expanded to the whole of the Mediterranean region.
5. OB JECTIVE OF IMPRO VEMENT
Main fig br eedingobjectives ar e (Storey, 1975; Jona and Gribaudo, 1991):
i. Development of high yielding cultivars.
ii. ImprovementinfruitqualityHighfruiteatingqualitywithimprovedstorageability.
iii. Eliminationofcaprification.
iv. Transfer of nematode and insect resistance characters of wild fig varieties to high
yieldinggoodqualitycultivars.
v. Persistence of syconia to ripeness.
vi. Resistance to pests and diseases.
vii. Hardiness to environment constraints.
6. CYTOGENETICS
The common fig (Ficus carica) is diploid, the somatic chromosome number is 2n = 2x
= 26 (Storey, 1975;Jona and Gribaudo, 1991) whileF.elasticacv. Decora is triploidin
nature. Ficus carica is gynodioecious, bearing either hermaphrodite or female fig on
separate plants. Males are the heterogametic (GA/ga) and female are the homogametic
(ga/ga) (Storey 1955). Shortstyle female flowers and male flowers are determined by a
pair of dominant genes on the samechromosome (GA); recessive alleles resultin long
style female flowers and suppression of male flowers (ga).
7. INHERITANCE OF PATTERN
Knowledgeof magnitude ofgenotypic and phenotypicvariation, genetic gainand genetic
advancearethemajortoolsforselectingefficientbreedingprogramme.Infig,quantitative
inheritance is a feature of many important traits, such as yield, quality and disease
resistance.Amongthedifferenttraitsthoseofhighdiscriminatinglevelwereleafdimensions,
shootdimensions,petiolecolor anddimensions,depthofsinus,fruitshapeandcolor,fruit
weight,dimensions,ostiolediameter,peelcolourandjuiceacidity.Amongthesecharacters,
some were good criteria for discriminating between cultivars. Several studies showed a
positive correlationbetween yield and fruit weight, fruit diameter, dry matter. A positive
correlation between polyphenols content and fruit weight and with flesh thickness; the
ostiole diameter with fleshthicknessandostiole opening arecommonlyobserved. Ostiole
sizehas interrela tionship with pests and diseases aslar ge ostiole an undesirable property
because pests and pathogens enter the fruit. Wide phenotypic and molecular diversity
found in fig germplasm indicates a considera ble potential for impr oving this crop.
152Breeding of Un derutilized F ruit Crops
8. PROBLEMS IN BREEDING
Figbreedingisacomplexbecauseofinvolvingtwotreemorphs(Caprifig andEdiblefig),
three floral forms (longstyled female, shortstyled female, and male flowers), and the
insect pollinator (Beck and Lord 1988).
i. Occurrence of Parthenocarpy and seedlessness.
ii. Complex pollinationmechanism i.e. caprification.
iii. Lack of reference collections.
iv. Lack of universal descriptor list to differentiate different varieties/species.
v. Presence of large intravariety diversity.
vi. Closeness of wild types to cultivated plants in some regions.
9. FLORAL BIOLOGY
9.1.FlowerBudInitiation
MorphologicalanddevelopmentalstudiesconductedbyDuric etal. (1992)duringdormancy
andgrowth oncv.Tenicabearing a single crop and cv. Petr ovaca bearing twocrops per
year reveal that three typesof buds occur on oneyearold shoots. (i) Flower buds which
arespherical,have35scales,withtheinflorescenceaxisintheinternalcavityenclosing
femaleprimordiadifferentiatedtovaryingdegreesdependingonthepositionofbudonthe
shoot.Thesebuds produce parthenocarpic fruits only in cv. Petrovaca. (ii) Mixed buds
are conical, madeup of 58 scales, and inflorescences on shoots ar ising fromthese buds
producefruitinbothcultivars.(iii)Vegetativebuds,madeupof35scales,whichusually
remain dormant and only rarely produce shoots..
9.2.Flowers
The flowers are three types.
i. male flowers with 15 stamens
ii. female flowers with ovary and long style, each resulting one seed
iii. gall flowerresemblesfemaleflower, short styled,ovary swollen, hiversfigwasp and
does not set seed.
Theflowersarehollowandbornintheleafaxils,reducedtoessentialorgansi.e.male
and female and are produced inside globosepear shaped receptacles with narrowmouth
(ostiole). Flowersarebornaxillaryonpreviousgrowthandalsooncurrentgrowth.Near
the ostiole male flowers are borne and towards the stalk female flower are present.
In Capri fig all three types of flower occur inside each fig whereas, in other types
(dioecious),themaleandgallflowersareproducedinsame inflorescenceononetreeand
female inflorescence on another tree.
Fig(Ficus carica)153
Fig is a monoecious, deciduous tree or large shrub. The subgenus Eusyce to which
Ficuscaricabelongsischaracterizedbyhavingunisexualflowers onlyandgynodioecism
(Storey, 1975). The inflorescence is unique, consisting of a syconium (the fig), which
enclosesmanyunisexualflowersthatcanbeaccessedviatheostiolebypollinatingwasps
that may give the true fruits. Tiny pedicellate druplets usually called the ‘seed’ (Storey,
1975; IBPGR, 1986). Morphologically Ficus carica is considered gynodioecious but
functionally dioecious. According to the sex of the flowers in the syconium, two main
types of trees can be distinguished. The first one is the caprifig in which the syconia
containshortstyledpistillateflowersdistributedovermostoftheinnerwallandstaminate
flowersaroundtheinterioroftheostiole. Theotheroneisfemalefigtreewiththesyconia
containingonlylongstyledpistillateflowers.Normally, onlythelateroneproducesedible
figs(Valdeyronand Lloyd, 1979; Storey, 1975).
Ficus carica is a gynodioecious species, having two distinct forms of trees; the
caprifig is monoecious, and the common fig is pistillate (Storey, 1975). Smyrna fig
produces male flower and proves to be unisexual from inception and shows ontogenetic
divergenceat primordial stage,whereasmale flowers areinitiatedas hermaphroditesand
undergo gynoecium abortion at the magaspore mother cell stage.A caprifig is seen as
expressing two pathways to unisexuality. Smyrna fig inflorescence produces abortive
hermaphrodite flowers at the same position as the male flowers in caprifig (Beck and
Lord,1988).
9.3.Pollination
Carles (1985) workingwith figin France reported that for ear ly fig production for fresh
consumption(whichismoreeconomic)pollinationistobeavoided.Pollinationisnecessary
forlatefigswhichareusedforprocessing.Valizadchetal.(1987)studiedthepollenflow
in4 femaleplants(domesticfig)and5functionallymale plants (caprifig)andfoundthat
eachsynconiumwaspollinatedtoabout80percentbyasingleinsectdespitebeingvisited
by several. The probability that a male plant pollinates a female plant was negatively
correlated with the distance between them, but at distance over 1020 m the corr elation
could be marked by competition between femaleplants. In fact, Smyrna type figs will
never set fruit without pollination from caprifrg.
Thepollination and development of flowersofthecaprifigandfemale figclone were
studiedbyTanriveretal.(1997)in Turkey.Thedevelopmentofthepistillateandstaminate
flowers was monitoredby stereomicroscope. Inthe female tree, the synconia of the main
crop were initiated on 1015 Ma rch; initiation of the pistillate flowers was observed on
22April.At theend ofMay, some figwasps (Blastophaga psenes) entered the synconia
andpollinatedthepistillateflowers.On21June,thestylehadturnedbrown,theovulewas
swollen andthe testa washardened. In the male trees, in early andmidApril, the wasps
released from the mammoni crop (initia ted on currentgrowth in the summer :matures in
autumn)enteredintothesynconia oftheprofichicrop(initiatedinspringfromlatentbuds
154Breeding of Un derutilized F ruit Crops
on the previous season’s wood; matures in summer) and laid their eggs into ovary. The
gall and male flowers were already mature (midAprila nd early June, r espectively when
the female flowers wereinitiated at the time when the wasps were ready to emerge and
enter the svnconia of the mamme crop (initiated in autumn on current season’s growth,
matures in spring) to layeggs. Thefragrance of the receptive fig attracts and stimula tes
fig pollinators (Blastophaga psenes) (Gibernau et al. 1998 ). Fig species emit specific
compoundsinordertoattractpollinatingwasps.Volatilecompoundsreleasedbyreceptive
figsfrom13tropicalFicusspecieshave been identified.Theyare mainlyterpenoids. but
also include benzenoids and nonterpenoid oxygenated compounds. Pollen is actually
transferred by a small wasp (Blastophaga psenes) which over winters in the pollen
producing caprifig( Gerdts andClark, 1979).
Thefigs commonlygrown inIndiaareparthenocarpic innatureanddonotneed any
crosspollination from wild fig (capri fig) which is very common in other fig growing
countries.However,fruitsettinghereisalsoinhibitedundercertainconditions.Ithasbeen
suggestedthatparthenocarpyisfavouredorinhibitedinagiventypebyclimaticcondition
of the placewhereit is growing. Thus, Pune,Black Ischia and Brown Turkey have been
found to be parthenocar pic a tKadur, while Turkish White hasfailed to set fruit without
caprification. Likewise, Pune and Black Ischia do not set fruits without caprification in
Allahabad.Infact,Smyrnatypefigswillneversetfruitswithoutpollinationfromcaprifig.
9.3.1.PollinationMechanism(capriûcation)
Fig has interestingpollination mechanism involving a symbiotic rela tionship between the
plant and its pollinator wasp Blastophaga psenes L. The ûg relies upon the wasp for
seed production, and the wasp complete life cycle within the inflorescence or syconium.
Female and male unisexual flowers are produced but female flowers mature before the
males. Within the syconium, the female wasp lays her eggs and pollinates the female
flowers. Therearetwotypesoffemaleflowerwithinthesyconium, shortstyled andlong
styled. T he wasp penetrates the shortstyled flowers with its ovipositor and lays an egg
intheovary. These shortstyledflowersbecomegallsasthedeveloping wasp larvae feed
onthe ovarytissue. Thestyle ofthe longstyledfloweris longer than theovipositor, and
these flowers are pollinated by thefemalewasp with pollen collected from a male stage
syconium. The syconia and itsseed then develop slowly a s the wasp larvae grow. When
thewasps, bothfemaleandmale,haveemergedfromtheirgallswithinthesyconium,the
male flowersof the syconium aremature. The wasps mate withinthe syconium, and the
males then die, having spent their whole life in this enclosure. Fertilised females collect
pollen from the male flowers, leave the male stage syconium and carry the pollen to a
female stage syconium, entering via the ostiole. This whole process is termed as
capriûcation.
The caprifigs (with wasp inside) are collected and placed in small bags or wire
basketsthatarehunginthefruitingSmyrnatypetrees.Theemergingwaspscoveredwith
Fig(Ficus carica)155
pollen,entertheSmyrnafruits andpollinatethelongstyledpistillate flowersinside (Galil
and Necman, 1977). In China,Yang et al. (1999) observed that fig is dependent on fig
wasps of the family Agaonidae for pollination and in turn the fig wasps are dependent
upon the galls (ovaries) of the figs for larval development and a complicated symbiotic
(mutualistic) relationship between the two has developed. Ramireza nd Malavasi (1998)
reportedthatpollinationofthepistillatefigflowers(Ficusspp.)hascrucialeffectsforthe
figs and the pollinating wasps. It allows normal development of seeds and wasps. Thus,
for fruit setting of the Symrna fig, caprifig trees as well as the Blastophaga wasps are
necessary. In addition to pollination, these fig wasps also carry propagules, mainly of
Fusarium verticillioides [formerly F. moniliforme (Gibberella fujikuroi)] and other
Fusarium spp. which causeendosepsis, from pollinizer figs tothe edible Calimyrna figs
inCalifornia.MichailidesandMorgan(1998)foundthatendosepsisdecreasedwithdistance
from the source, decreasing faster to the south than in other dir ections from the source.
Thefragranceofthereceptivefigattracts andstimulatesfigpollinators(Blastophaga
psenes) (Gibernau et al. 1998 ). Fig speciesemit specific compounds in or der to attra ct
pollinating wasps. Volatile compounds released by receptive figs from 13 tropical Ficus
species havebeenidentified.Theyaremainlyterpenoids. but alsoincludebenzenoidsand
nonterpenoid oxygenated compounds. Pollen is actually transferred by a small wasp
(Blastophaga psenes) which over winters in the pollen producing caprifig ( Gerdts and
Clark, 1979). Thus, for fruit setting of the Symrna fig, caprifig trees as well as the
Blastophagawaspsarenecessary.However,caprificationisacumbersomeandexpensive
process.
Considerablesuccesshasbeenachievedbysubstitutingplantgrowthregulatorsprays
for caprification. Of the numerous substances tested, IBA, NAA, 2,4,5T and 4CPA
proved effective in inducing early maturing parthenocarpic figs (Crane and Blondcau,
1949,1950).Theparthenocarpicfruitsdeveloptoa normalsizeandhavedesirablesugar
content, but as theyarecompletelyseedless,thebaking industries,whichusemostofthe
CalimyrnafigsproducedintheUSA,donotwanttouseit,becausetheylackthecrunchy
quality imparted by fig seeds. Subsequently, Crane (1952) found that BOA induced
parthenocarpyandtheformationofdrupeletswithhollow,sclerifiedendocarp,butitisstill
not acceptable to the industry.
9.4.Fruit
Theediblefigisamultiplefruit.Botanicallyitisknownassyconiumwhichconsistsofa
fleshyhollowreceptaclewithanarrowapertureatthetipandnumeroussmallflowerslining
theinnersurface.Thetruefruitsarethetinydrupeletsinsidethecavityofthefusedpeduncle.
Thefigisanaggregatefruitcomposedofindividualsmalldrupes;eachistermedadrupelets.
Thefigfruitsareborneintheaxilsoftheleaves;twoinflorescencesandonevegetativebud
arepresentatthesamelateralpositionintheleafaxils.Butincultivarssuchas‘Mission’and
‘BrownTurkey,’usuallyonlyoneinflorescencedevelopsintoasyconium,whilein‘Kadota’
156Breeding of Un derutilized F ruit Crops
and ‘Calimyrna’ cultivars , often both inflorescences at a node may develop. Fig fruit
development follow double sigmoid curve with three defined growth periods (Crane and
Brown1950;CraneandBaker1953). DuringIIphaseofgrowth,about70% ofthetotaldry
weightand90%ofthetotalsugarcontentisaccumulatedinthefruit.
9.4.1.BearingBehaviour
Thefigmaystartbearingafewfruitsayearortwoafterplanting,butthiscropisnotallowed
todevelop,becausethecropmayreducethevegetativegrowthoftheplant.Usually,steady
yieldcanbeobtainedfromthefifthyearonwards.Thetreescontinuetobeargoodcropsfor
3040 years before they begin to decline. The fruits start ripening from MarchMay in
westernIndiaandMayJuneinnorthernIndia.InSouthIndia,thefigbearstwiceayear
onceinJulySeptemberandagaininFebruaryMay.ThefigscommonlygrowninIndiaare
parthenocarpicinnatureanddonotneedanycrosspollinationwithwildfig(caprifig),which
isaverycommonpracticeinothercountries.However,fruitsettingisalsoinhibitedunder
certainconditions. It has been suggested that parthenocarpyis favouredor inhibited in a
giventypebyclimaticconditionsoftheplacewhereitisgrowing.Thus,Pune,BlackIschia
andBrownTurkeyhavebeenfoundtobeparthenocarpicatKodur,whileTurkishWhitehas
failedtosetfruitswithoutcaprification(Naik,1949).Likewise,PuneandBlackIschiadonot
setfruitswithoutcaprificationinAllahabad(Hayes,1957).
9.4.2.Fruit Growthand Development
Fig is a gynodioecious species and some female type’s need pollination while others set
fruits parthenocarpically. Pollination is effected by a wasp, which develops inside the
syconium of male fig. This symbiotic relationship is a classical case of coevolution
betweena plantandinsect. Figfruits show adouble sigmoid growthcurvetwoperiods
ofrapidgrowthbeingseparatedbyaperiodofslowgrowth(Crane,1948).Inaninvestigation
on fruit ripening in cv. Rampelina, samples were collected at four fruitgrowth stages
duringJune,July,August andSeptember anddividedby weight into6groups:05,610,
1115,1620, 2130and>30 g.
Fruitdevelopmentfollowedadoublesigmoidpattern.Respirationandethyleneproduction
decreasedsharplyduringcellmultiplicationinthefirstgrowthperiod,remainedconstantinthe
stasisperiodandthenincreasedrapidlypriortotheonsetofthesecondgrowthperioduptothe
climacteric. Respirationdecreasedsharplyinthesmallestfruits,whereasitincreasedinthe
largest fruits during midAugust in conjunction with the onset of the second rapid growth
period.Ethyleneproductionwasinitiallyhighinthesmallestfruitsbutdecreasedprogressively
with the onset of maturity; only fruits >30 g showed an incr ease in ethylene production
concomitantwiththeonset ofripening.Reducingsugarcontentremainedlowthroughoutin
smallerfruitsbutincreasedwithripeninginthelargestfruits(Chessaelal.,1992).
Growthofsyconiumin8cultivarswasstudiedbyElKassasetal.(1992c).Therapid
growthcycleperiodIlasted5to6weeksinallcultivars.However,periodIIvariedsubstantially
Fig(Ficus carica)157
amongcultivarsandlasted8weeksinSultani,5weeks inAbiadAsswan,Kahramaniand
BrownTurkeyand34weeksinConadria,Abboudi,DiRedoandBlackMission.Thethird
growthphasewascharacterizedbyanacceleratedincreaseindiameterofsyconiauntilfruit
wasripe.PeriodIIIlasted5weeksinBrownTurkey,4weeksinConadria,BlackMission
and DiRedo and 3 weeks in Sultani.Overall, syconia developed over a period of 1216
weeks.IthasbeenknownsincethethirdcenturyBCthatgrowthandmaturationofthefig
fruitcanbeadvancedbyafewdaysifadropofoliveoilisappliedtotheostiole(eye)during
thetendayperiodfollowingthetimeatwhichalldrupeletsinthefruithaveturnedred(Sand
et al., 1969). T his period brackets the transition from stage two to stage three of fruit
growth. The stimulating agentis ethylene, which is produced as a breakdown productof
oliveoil,especiallywhentheoilisexposedtosolarradiation.
It has been found that application of ethylene exogenously in the first growth phase
usuallyretardsfruitgrowth,butacceleratesdevelopmentandripeningwhenappliedinthe
secondgrowthphase,thefruitsareripenwithinaweekafterapplication(MareiandCrane,
1971;Puechetal., 1976).Duringthesecond periodofsyconinmgrowthof2successive
seasons,shootsoffigcv.Sultanitreesweresubjectedtooneofthefollowingtreatments:a
singlesprayof100,200or300ppmethephon.girdling,manualremovalofthe5leastmature
fruitsattheendoftheshoot (thinning), girdling+ethephon,manualthinning+ethephon.In
bothseasons,alltreatmentsexceptmanualthinningsignificantlyhastenedfruitmaturityand
improvedquality,comparedwithuntreatedcontrols.Themosteffectivetreatmentwas300
ppmethephoncombinedwithshootgirdling.Applicationof500ppm ethrelresultedin82%
ofthefruitsbeingharvestedat thefirstof3harvestscomparedwith56%fromuntreated
trees.Damagecausedbyinsectsorfungalinfectionwasless(Fergusonetal.,1990).
The typical growth curve of the Bursa Siyahi fig fruit was shown to be a double
sigmoidwiththelengthofgrowthperiodsI,IIandIII being44,35and13days,respectively
(Celikel et al., 1998). Ethrel (ethephon) at 100, 250, 500 or 1000 ppm was sprayed at
intervals during growth phase II, on the leaves and fruits of the cultivar Bursa Sivahi.
Ethrel at 250 or 500 ppm sprayed at a late stage of phase II stimulated growth and
shortenedthetimeto maturitywithout affecting fruit quality.Ethreltreated fruits ripened
5 days earlier than the contr ol. Ethylene treated fruits attain normal size, colour, texture
andflavourmuchearlierthantheuntreatedones;theyalsoshowahigher contentoftotal
soluble solids and dry weight (BenYehoshua et al., 1970; Gerdts and Obenanf, 1972;
Mougheith and ElBanna. 1974).
Rapeseed oil injected through the ostiole of fig cv. MasuiDauphin fruits harvested
beforetheclimactericinducedrapidethyleneevolution,similartothatobtainedbytreating
with exogenous ethylene (Gao et al., 1991). Ethylene.forming enzymc (EFE) activity
increased rapidly, reaching a level 12 times higher than the controla fter 24 h, but ACC
contentincreasedslowly.Oilinducedethyleneproductionwasalmostcompletelyinhibited
by aminooxyacetic acid (AOA) an inhibitor of ACC svnthase. Apart from the basic
essentialoil,2ethyl1,2dihydrothiophene,theripefruitshavevolatilephenols,inparticular
vanillinoheterocvcliccompounds,acetone,aliphaticacids,terpenealcohols,aliphaticand
aromatic alcohols and aldehydes (Pisarnickii et al., 1986).
158Breeding of Un derutilized F ruit Crops
The quality breba fruits of figs are affected by plant nutrition in addition to cultivar
characteristicsand ecological conditionsduring ripening (Sahinet al., 1999). Irget et al.
(1999) reported that fruits from trees given Ca(NO3,)2, were harder in texture when
dried than those from trees given KNO3. Ferrara (1999) reported that bagged fruits
showed faster growth and could be harvested 68 days before unbagged controls in the
same area and 2030 days before inland and hill orchards.
10. DIFFERENT SPECIES
ThegenusFicus,whichis comprised ofabout 700 species,classifiedintosix subgenera,
characterized byaparticular reproductive system(Berg,2003). It is nativeto theMiddle
East particularly southern Arabia where wild and caprifig trees are still to be found
(Condit,1947;Zukovskij,1950;Storey,1975).
Ficuscarica (Co mmon fig)
It is a small to moderatesized deciduous tree, 68 m high with a short twisted trunk,
crown with irregular branches; frequently shoots develop at the base of the trunk. The
bark is pale grey. Terminal buds are short and stout. Leaves are broa d, ovate or nearly
orbicular,moreorlessdeeply 35lobed,roughaboveandpubescent below;longstalked,
leaf blade 1025 cm long, dark green with pronounced venation. The female fig plants
havelargerleavesanddenser,morespreadingcrownsthanmales.Leaffallbeginsearlier
inmales. Theleaves and the spongy parenchyma tissues are thicker infemales. The leaf
thicknessis125.5µminfemalesand111.8µminmales(Kotaevaetal.,1982).Fruitsare
mostly solitary, axillary, green or yellow, brown, purplish or even black depending on
cultivar, more or less pearshaped with either a velvety or glabrous skin, in certain
cultivars upto 6cm in diameter, but normally of moderate size.
11. TYPES AND CULTIVARS
Depending upon the nature of flowers and the method of pollination, there are four
pomologically distinct classes of fig: (i) Common fig or Adriatic fig, (ii) Capri fig (iii)
Smyrna fig and (iv) SanPedro fig.
11.1. Common Fig
Inthistypetheindividualflowersarepistillateandthefruitsdevelopwithoutthestimulation
ofpollinationandfertilization. SomecultivarsofthistypeareKadota(Dottato),Mission,
Adriatic, Brown Turkey, Celeste and Conadria.
11.2. Capri Fig
It is themost primitive cultivated type with short st yled pistilla te flowers andfunctional
staminate flowers. Most caprifig are not edible but are grown because they harbour a
small wasp, Blastophaga psens which is necessary for pollination and setting of fruits.
Fig(Ficus carica)159
11.3.Smyrna Fig
Inthistype,thefruitsdonotdevelopunlesstheflowersarepollinatedwithpollencarried
from the male flowers ofthe caprifig by the tiny Blastophaga wasp. Calimyrna is the
most common and widely grown cultivar of Symrna type.
11.4. San Pe dro Fig
Itisanintermediatetypeinwhichthefirstcropknownas‘breba’.Thecropiscompletely
parthenocarpicanddoesnotrequirepollinationandfertilizationofflowers,butthesecond
cropdevelopsonly iftheflowersarepollinatedasinthecaseofSmyrnatype.SanPedro,
King and Gentile are common cultivars of this type.
Four types of figs are described basedon cropping and pollination characteristics.
Table. Horticul tural classifica tion of fig
Type Popul ar varieties Flower Mode of No.of Ot her feat ures
type pollination Crops
EdibleFig Poon a,K adota, Lon g styled Fru its develop s 12 So me variet ies
Con ardia , Missio n, pi stil late par then ocar pical ly pro duce a sm all
Brown Turkey flo we rs with out p ollinat ion breba or firstcr op
Smyrna Ca limyrna (S arilop ), L ong styled F emal e wa sps 1 In a ddition to main
Zidi, Taranim t pis till ate emerg ing f rom th e or second c rop
flo we rs spr ing ca prifig orig inated fr om
enter smyrna fig the caprifig. T he
for o viposition & fer tile seed s
in the pro cess of contribute to t he
po llin atio n excellent fruit
quality
Sanped ro King, G entile, San L ong styled First c rop (b reba) 2 Com merc ially no t
Pedro , Dauph ine, pi stil late d evelops with out impo rtant s ome
La mp eir ia flo wer s pollination but the white large fruited
main (seco nd) types are grown in
cro p requ ires med iter rane an
po llin ation c ou ntrie s
Wildfig/ Road in3, S amson, sh ort styled S elf fertile 3 A primitive type.
caprifig Stanford ,Brawley pis till ate ( pers isten t) Fru its have almost
(male/goat flow ers & syc oni a no edible value,
fig) fu nctio nal pro vides the
st ami na te sourc e of po llen
flow ers nea r for c omm ercial
the o stiole p lanting s of
cau ducou s figs.
(Saleeb and Storey 1 975)
160Breeding of Un derutilized F ruit Crops
Of these four types, Smyrna fig is commercially most important and is extensively
grown in Europe a nd USA. The cultivars Kadota. Krymskii 43 and Bianco Grossowere
recommended for cultivation in Southern Uzbekistan (Mysina, 1985). It has a superior
nutty flavour due to presence of fertile seeds. In India, common fig is mostly grown. It
is considered tobe a hybrid between imported F. caricaa nd indigenous species. A large
number of cultivated forms are grown in which the fruits vary in shape, size, colour of
skin, colour and flavour of flesh and period of ripening. Someof the cultivars grown in
India are Black Ischia, Brown Turkey. Turkish White, Kabul and Marseilles. The figs
growninmanypartsofIndiaarenamedafterlocality,buttheydonotexhibitanyspecial
distinctionthat warrantsvarietalnames.Punefigisofmediumsize,bellshapedandlight
purpleincolourwitharosyflesh.In SouthIndia,PunefigandMarseillesarecommonly
grownthelatterthrivingwellinthehillyregions.ThefruitsofMarseillesaremediumsize,
palegreen on the rind with a whitish sweet flesh. Condit (1955) gave a good a ccount of
711 cultivars of figs and listed 946 synonyms in addition to 98 which hementioned as
having been described by other workers.
Lauri et al. (1993) classified edible fig cultivars into three horticultural categories:
Smyrna, San Pedro and Common. San Pedro and Smyrna types require caprification
(unlike thecommon type), thefirst produces breba andmain (or second) crops, whereas
the second produces only main crops. Morphological observations on cultivar s Grise de
Tarascon(brebaandmain)andRougedeBordeauxor Pastiliera (mainonly)duringtheir
fourthyearofgrowthshowedthatbrebaandmaincropshavedistinctdistributionsalong
the annual gradient;br eba s are located onthe neoformed part of shoot axes, while main
cropfigsareonthepreformedpart.Thisworkinvestigatedrelationshipsbetweenvegetative
growthandspatialdistributionofsyconiaalongashootaxis,anddistributionofaxeswithin
the crown as determined by architectural analysis.
Fig.Floralmorphologyincaprifigs and edible figs. (Armstrong, 2006)
Fig(Ficus carica)161
12. CROP IMPROVE MENT METHODS
Althoughfighasbeenundercultivationformanycenturiesandisnowdistributedinmany
countries of the world, relatively little work has been done for its improvement. The
earliestreportsonfigbreedingarethoseofSwingle(1912),Hunt(1912),Burbank(1914),
andCondit(1928).A projectentitled‘Genetics,cytology,morphologyandbreedingoffigs
(Ficuscarica)isbeingoperatedat theUniversityofCaliforniasince1922andasaresult
ofcontinuousplannedbreedingworkelevennewAmericancultivarshavebeendeveloped.
12.1.Selection
Althoughfighasbeenundercultivationformanycenturiesandisnowdistributedinmany
countries of the world, relatively little work has been done for its improvement. Most of
the fig cultivars cultivated today are selections made by unknownpersons inAsia and
Europe in the past centuries from wild seedling trees andchance seedlings. Since then
these have been maintained clonally by rooted cuttings and in course of time, have
acquired names. Three fig cultivars were selected by Ozeker and Isfendiyaroglu (1998)
inthe Ciftlikkoy region inTurkey(C1, C2 and C4)fr om 12 figcultivars that bear ear ly
breba fruits. The average fruit weight ranged between 3090 g, the total soluble solids
content in fruit juices was between 16 and27.6% titrable acidity as citric acid between
0.06and0.15g/ml,pHvalue between4.735.90andthefruitfleshfirmnesswasbetween
0.201.20 kg/cm
2
. In Italy, Grassi (1998) identified 442 clonesfrom different regions.
Morphological, phenological, productive and carpological observations cleared up many
synonymiesand led to thegroupingof manycloneswith similar characteristicsinto only
50 groups ofdifferent values in cultivation.
Chenetal.(1997)evaluated21cultivarsovera yearperiodforgrowthofshootsand
fruits,freezing injuryandothercharacteristicsinXuchangareainChina.Thebestvarieties
forHenanprovince wereMasuiDauphine, Brunswick,He18, andentries 8 and10. The
freezinginjurycouldbealleviatedbytrainingofbranchesintoacompactform.InTunisia,
there is a large varietal diversity with some varieties being threatened by extinction.
Development of successful modern production would depend on varietal selection and
improvement combined with resolution of some of the problems currently plaguing the
industryfruit cracking,handlingproblems environmentalimpact.
12.2.Hybridization
The classical breeding approaches in fig were summarized by Condit (1947), Storey
(1975),Obenaufetal. (1978), Fergusonetal.(1990), andMars (2003). Fig breeding is
a complex because of involving two tree morphs (Caprifig and Edible fig), three floral
forms(longstyledfemale,shortstyledfemale,andmaleflowers),andtheinsectpollinator
(Beck and Lord 1988). Two breeding progr ams were conducted in the beginning of the
20thcentury attheUniversityof California,led by I. J.Condit andN. B. Storeyandby
162Breeding of Un derutilized F ruit Crops
E. N. O’Rourke Jr at Louisiana State University. In addition, fig crosses and their
evaluation has beendescribedin Japan (Awamiir a, et al., 1996). Hybridiza tion began in
2003,andbytheendof2006,about4,000hybridfigseedlingswereplantedattheVolcani
Center,BetDagan,Israel.TheU.S.breedingprograminDavis,CaliforniabyConditand
Storeyproduced 300hybridprogeniestotalingmorethan30,000seedlings(Storey1975).
ThemostsignificantachievementoftheCaliforniafigbreedingprogrammewasdevelopment
offivehybridcultivars:‘Conadria’,DiRedo’,‘Flanders’,‘Tena’,and‘Excel’.Morerecently
Doyle and Ferguson(2005) released the cultivar‘Sierra’, a commonfig typewith green
fig color and late summer maturity.
12.2.1.BreedingTechnique
Storey (1975) described in details the breeding technique being followed in fig. Trabut
(1922) and Condit (1947, 1950) tried interspecific hybridization involving F. carica, F.
palmata and F pseudocarica. However, the hybrids thus obtained were not of much
commercial value. In the Badkhyz area of Turkmenistan (former USSR) where F.
afghanistanica coexists with F. carica, spontaneous interspecific hybrids have been
found which are intermediate between parents (Petrova and Popov, 1979). New early
flowering caprifigswere bred at the Nikita Botanical Garden in Cr imea (former USSR)
using seedlings obtained by intervarietal and interspecific hybridization. Twelve new
caprifigs were selected which arc early flowering and produce an abundance of viable
pollen(Arendt andAleksandrova,1971).Arendt (1974) reviewedthebreedingworkdone
in former USSR. She obtained best results by interspecific hybridization using a hardy
triploid species referred to as the Afghanistan fig (F afghanistanica) and F. carica
Yellow.T he most significant achievement in the history offig breeding, however, comes
from the researches done in California. The fruit characters of eleven new hybrids
released and adapted for commercial cultivation in California.
Toclarifytheeffectofvariousseedparentsonfrequencydistributionofparthenocarpic
characteristicamongprogeniesoffig(Ficuscarica),selectedfigcultivarswerepollinated
with pollen from incomplete parthenocarpic caprifig cv. Caprifig 6085.After separating
the seedlings into a caprifig or a common fig type, the occurrence of parthenocarpy in
the first or second crops was investigated. In the first crop of caprifig type, a high
frequencyofparthenocarpywasfoundinprogeniesfromSanPedrotypecultivar,whereas
thecommontypecultivarseta moderatenumberofparthenocarpicfruitinthefirstcrop.
In the second crop, thecommontypecultivar producedprogenies witha highfrequency
of parthenocarpic fruit. Almost no parthenocarpic seedlings were producedwhen a non
parthenoca rpicSmyrnatypecultivar was useda s a seed parent (Awamura et at., 1996).
Fruit characters, fruit weight (FW) and soluble solids content (SSC) in the juice of
hybrids of 9 cross combinations were investigated byAwamura et al. (1997). Family
means of the groups, namely caprifigtype first crop, figtype first crop and figtype
second crop, were determined. There were positive correlations between the FW of the
Fig(Ficus carica)163
3groupsandmidparentalvalues,andbetween theSSCoffigtypefirstandsecondcrop,
and midparental values. There were negative correlations between the FW of caprifig
typeandfigtypefirstcropandthemidparentalSSCvalue,andbetweentheSSCoffig
type second crop and midparental FW value.
O’Rourke (1966), Puls et al. (1967) and Puls and O’Rourke (1967) have reported
significant progressin breeding figs for rootknot nematode (Meloidogyneincognita var.
acrita) resistance in Louisiana.Afairly highorder of resistance was found in individual
seedlings in progenies of the commercially grown cultivars Hunt and Celeste.
CultivarLSUwasdevelopedfromthepollinationofHuntbyanunknownCalifornian
caprifig designatedas Cl. LSU Purple is parthenocarpic, so thereare few seeds or kerfs
inthepulp. Theflavourisgood andmildandsugarcontentishigh. Thetreeisvigorous,
upright,aboveaverageforcoldtoleranceandresistanttoleafdiseasesandrootnematodes
(O’Rourke, 1992). Diploid apogamy was induced in F. carica and F. afghanistanica by
application of Lilium candidum pollen and various physiologically active substances.
Adenosine triphosphate stimulated apomictic seed development in F. carica, as did
naphthylacetamide in F. afghanistanica (Romanova, 1979). Neeman and Galil (1978)
showed that artificial pollination of bagged spring and summer caprifig syconia resulted
in seed set and sweet figs were obtained.
12.3.MutationalBreeding
Inducedmutationsmayplayanimportantroleinfigimprovementforimportanttraitssuch
assmallostiolsize,largefruitsize,fruitfleshquality,andtreeproductivityasconsiderable
diversity for fruit shape, color and tree vigor exist in many clonally derived cultivars
(Santoniet al., 2000). Mutational variation canbe induced either by specific treatments
withphysicalandchemicalmutagens orbytissueculture.Theeffectof gammaradiation
on cuttings, seeds, and pollen in figs has been well reported. The frequent finding was
dwarfness andaccelerationoffruiting (Mars 2003). Somemutantswereusedinbreeding
programsandthecultivarBol’obtainedbyseedirradiationfollowedbyselection(Akhuid
Zade1981).Inaddition,spontaneousmutantscanoccurincommercialorchards.Recently
‘RedKadota’,amutantof‘Kadota’selectedfromcommercialorchardinIsraelissimilar
to ‘Kadota’ cultivar, but has red skin that develops during fruit maturation; therefore, it
was named and registered as ‘Red Kadota’ (Flaishman et al., 2007). In vitro tissue
culture methods may also be very useful in selecting new cultivars from somaclonal
variation(Mur iithi et al., 1982; Pontikis and Melas 1986), but none have been r eleased
yet from tissueculture mutagenesis (Mars 2003).
12.4.Biotechnology
12.4.1.MolecularBreeding
Compared to traditional breeding techniques, application of genetic engineering to
homologous and/or heterologous genetic material into fruit trees, offers the potential of
164Breeding of Un derutilized F ruit Crops
obtainingimprovedplantingstocksinashortperiodoftime. Efficienttransformationcan
also be used for the production of heterologous polypeptides having nutritional and/or
pharmaceutical value.Yakushijiet al.(2003),reportedadventitious shootregenerationin
tissuecultureinF.carica.Theyalsoreportedamethodfortheinductionoforganogenesis
fromleafexplantsofF.caricausingphloroglucinol(PC).Yancheva etal.(2005),reported
an efficient and reproducible system for regeneration of ‘Brown Turkey’ and ‘Smyrna’
fig.Inaddition, efficient andreproducible transformationsystems for bothcultivarswere
alsoreportedbyYanchevaetal.(2005).Similartoregeneration,theorientationoftheleaf
surfaceduringorganogenesiswasakeyfactorforsuccessfultransformation.Leafexplants
of invitro propagated plantswere cocultivated with the disarmed Agrobacterium strain
EHAI05harboringtheplasmidpME504thatcarriedtheuidAintron,barandnptllgenes.
Transformants were obtained by selection on the antibiotic ‘I Kanamycin at variable
transformation efficiencies in different cultivars (Yancheva et al., 2005). The transgenic
nature of the regenerated plants was confirmed by PCR and Southern blot and gave
typicalstainingforthereportinggeneGUS.Histochemicallocalizationof13glucuronidase
(GUS)activityconfirmedthatthecauliflowermosaicvirus(CaMv)promoterfunctionsin
cells of the fig syconium.
12.4.2.MicroPropagation
Micropropagation of fig is also possible. Pontikis andMelas (1986) developed a micro
propagationtechniqueinwhichshoottipsofcv.KalamonwereculturedonMurashigeand
Skoog’s medium containing several growth regulators. Shoot proliferation occurred in
mediacontainingphloroglucinoat89mg/l.Theshootswerethenrootedinmediacontaining
IBA or IBA and NAA. The best rooting (80%) was obtained withIBA at 1 mg/l. The
rooted plants weresuccessfullytransferredto sterilevermiculiteandafteracclimatization
to pots containing equa l parts of peat and vermiculite.
A procedure formultipleshoot induction andplantlet regenerationwas developedby
Kumar et al. (1998) with apical buds collected from 7–to 8–year–old trees of Ficus
carica using MS medium supplemented with 2. 0 mg GA and 0.2 mg NAA/1. The in
vitroregenerated shoots were further multiplied onMS medium supplemented with 2.0
mg BA and 0.2 mg NAA/1; an average multiplication rate was four per subculture.
Excised shootswererootedinliquidhalfstrengthMS mediumsupplementedwith2.0mg
IBA/l and 0.2% activated charcoal.
ApicalshoottipsofthefigcultivarsBerberaandLampawereculturedbyNobreand
Romano(1998)onMuriithiorJonardculturemediumsupplementedwiththeantioxidants
likepolyvinylpyrrolidonc[(polyvidone)](PVP40)(0.025,0.05,0.1,1.0%v/v)orascorbic
acid (56.8 or 113.6
m
M). Growth and development were best on Muriithi medium
supplementedwith 0.05%PVP.Singlenodeexplantsofshootsobtainedintheestablishment
phase were cultured on basal MS medium supplemented with 3% sucrose and BA (0.7,
1.1 or 2.2
m
M) alone or in combinationwith NAA (1.0
m
M).After 3 weeks, the highest
Fig(Ficus carica)165
multiplicationrate(5.3shoots/culture)wasobtainedinthemediumsupplementedwith2.2
m
MBAwithoutNAA.Plantletsweresuccess–fullyacclimatized. Micropropagatedplants
produced fruits within 2 years after field–establishment. Apart from standardization of
protocol for micropropagation, biochemicalandmolecularmarkershavebeenusedinthe
identification of genotypes of fig.
12.4.3.BiochemicalandMolecularMarkers
Characterization of fig germplasms has been conducted by means of isoenzyme
electrophoresis on horizontal starch gel applied on cork tissue (Chessa et al., 1998).
Seventeen enzymes systems were analyzed and acid phosphatase (AcPH), diaphorasc
(DIA),fumarase(FUM),glutamateoxalacetatetransaminase(aspartateaminotransferase)
(GOT), malate dehydrogenase (MDH), peroxidase (PRX) and phosphoglucoisomerase
[glucose6phosphate isomerase] (PGI) showed high resolution of the bands and
reproducibility of the analysis. Only FUM had a monomorphicpattern, while the other
enzymes tested allowed the characterization of almost all fig varieties. The degree of
electrophoretic similarity between varieties was determined by calculating a similarity
index value based on three enzyme systems : AcPH, GOT and MDH.
Elisiario etal.(1998)usedisoenzymeandRAPDmarkerstodifferentiate55traditional
varieties of fig in Portugal. Six isoenzyme systems were revealed after starch gel
electrophoresisofleafextracts:glucose6phosphateisomerase(PGI),phosphoglucomutase
(PGM), isocitrate dchydrogcnase (IDH), malate dehydrogenase (MDH), aspartate
aminotransferase (GOT) and cytosol aminopeptidase (LAP). Three isozyme systems,
PGM, IDH and GOT revealed four polymorphic loci and were used for variety
characterization. Several clonesshared identical isoenzyrne banding patterns and could
not be distinguished by these markers. RAPDs were used in a second step of molecular
characterization. Sixty decamer primers were tested. Fortythree primers generated
amplifiedproductsandwereusedtodistinguishamongalltheclones,eitherwithingroups
of identical or different isozyme patterns. Similarly, identification of 21 fig accessions
(Ficus carica) representingdifferent var ieties wa sperformed using the RAPD technique
(Khadari et al., 1995). The 19 RAPD markers used provided 17 different banding
patterns.TheRAPDmarkersshowedsufficientpolymorphismforgenotypediscrimination,
clonalstability,environmentalstabilityandexperimentalreproducibility.
12.4.4.MarkerAssistedSelection
Markerassistedselectionormarkeraidedselection(MAS)isaprocesswherebyamarker
(morphological, biochemical or one based on DNA/RNA variation) is used for indirect
selectionof a genetic determinant or determinantsof a trait of interest(e.g. productivity,
disease resistance, abiotic stress tolerance, and/or quality). Yield, quality, and disease
resistance etc are important quantitative inheritance. Means of analyzing quantitative
variation and especially of uncovering its potential genetic basis are therefore of major
166Breeding of Un derutilized F ruit Crops
importa nce for breeding purposes.Many studieshave been interested in the development
ofothermethodsforcultivaridentification.Forinstance,proteinpolymorphismsincommon
figs were suitable for characterizing fig varieties. However, the success of varietal
identification depends on the number of isoenzymatic systems and alleles studied.
Unfortunately, this number is often limited, which leads to insufficient polymorphism
among closely related genotypes. Various DNA profiling methods ar e curr ently available
and consist of two ma in categoriesa ccording to the information provided: specificlocus
and codominant markers and arbitrary and dominant such quantitative variation results
from the combined action of multiple segregating genes and environmental factors. The
jointanalysisofgenotypemarkersegregationandphenotypicvaluesofindividualsorlines
enables the detection andlocation of loci affecting quantitativetr ait loci (QTL). In a fig
breedingprogram,QTLcanbeusedthroughtheapplicationofmolecularmarkers,providing
basisforsocalledmarkerassistedselection(MAS).Molecularidentificationoffigcultivars
has been carried out using isozyme markers (Cabrita et al., 2001), random amplified
polymorphicDNA(RAPD)(Cabritaetal.,2001;Galderisietal.,1999;Papadopoulouet
al., 2002), Mitochondrial DNA (mtDNA) Restricted Fragment Length Polymorphism
(RFLP)(Khadari etal.,2005)oramplifiedfragmentlengthpolymorphism(AFLP)(Cabrita
etal.2001).Khadari etal.(2001)identified8microsatellitesinfig.Morerecently,Giraldo
et al. (2005), reported 26 additional microsatellites that were tested in a group of 15
known fig cultivars. T hese tools canbe veryuseful in future evaluation of fig progenies
in fig breeding programs.
13. VARIETAL WEALTH
Therearemanycultivated varietiesineachclassoffigs.Infact, amongseveralhundreds
ofdescribedcultivars,75 %wereof‘Commontype’,18%of‘Smyrnatype’andtherest
(7 %)ar eof ‘San Pedr otype’ or capr ifigs (IBPGR, 1986). Over 700 varietal names are
in use but many are synonyms. Poona fig is most popular cultivar grown in India.
Bangalore,Bellary,Coimbatore,Daulatahad,Dindigual,Ganjam,Hindupur,Lucknowand
Saharanpur,haveclearlyacquiredthenamefromthelocationinwhichtheyarecultivated.
Mostofthemresembleinplantandfruitmorphologyto thatofPoonafig.Possiblythese
are either clones or ecotypes. Dinker,a nimprovement over Daultabad for yield andfr uit
quality,isgainingcommercialsignificance.Somewell known fighybrids fromCalifornia
haveperformed well in India in comparison to Poona fig. Excela nd C onardia figs that
developsmallercanopiesaresuitableforhighdensityplantingandfreefromfruitsplitting.
Conardia,ExcelandDeannaaregoodfordrying,canningandtablepurposes,respectively.
Table: Compre hensive list o ffig va rieties
Smyrnatype figs with s kin green B orsamele, Eisen, Kaab el Ghazal, Khazou ri,Ma la ki, Rosa,
or yellow; pulp white, amber, Sari Lop ( Calimyrna,Aidin), Sc ionto Sno wden. Sultanie
orverylightred
[Table Contd.
Fig(Ficus carica)167
Smyrnatype figs with s kin green Ab ate, Abi arou s, Ab ou che rch aou , Akc a, A lekak e, A mesa s,
or yellow ; pulp various s hades A r an im A m ell al , B ar d aj ic B lo we r s, C as t elh a no B ra nc o ,
ofred Chan gelge, Che faki, Chek er Injir, Ch oer, Cueritesto , Djafer i,
Dj ebali, Fiet ta, Hilg ard ,Isly, J adi, Kala mata , Karayap rak,
Kas saba, Kh adir, K ouffi Ve rt, L ebi, Mado ui, Malak i Blan c,
Mama ri, Mapl eLeaved Merchin i, Mot a , Mou zai, Pan ettaro ,
Pa sulito , Pazz o, Rixfor d, S esso, Si gilli, S oua bae lAdj ia,
Bl anc he, Stan for d, S ulta ne B ilo ng ue, Sultani , Tabelou t,
Tabelout, Tadefo uit, Taharit, Takourc hit, Tameriout , Taranimt,
Tau risano,Tazarift, Trêsum Prato, Verdescone, West, W ilson,
Yediver
Smyrnatype figs with skin dark Eusc aire, H amriti, Sc anc aniso
(various sh ades of violet to
black); pulp white or amber
Smyrnatype Figs with Skin Da rk A bo ug an djo ur, Ad jaff ar, A gh an, Ag ou arzg uil ef, A go us sim,
(VariousS hades of Red, Brown, Aranim Aber kane ,Ave rane, Avou zegar, Azen dja r, Barda kjik
or Vio let toBlack);Pulp Various Black , Bar li, Belman dil, Bes oulelKhadem, Bid dinel Brel,
Shades of Red Bo uAng , BouHar rak, Castelha no Pret o, Hab tir, Kara Injir,
Kha louli, Ko uffi Rou ge, L ugliatico , Marab out, Mo r, Mo r Güz,
Na polet ana, Passa can ne, P urp le Bullet in Sm yrna, Pu rple
Sm yrna, R her abi, Sala tello , So ltan ine, Tak lit, Tar ou mant ,
Tec hich, Turco, Zafrani
San Pedro typ ef ig s wit h skin Cach opeiro, Bra nco, Co lombro, Gen tile, Khdari, K in g, Noce,
green; pu lp red Pietri,San Giovanni, San Pedro, San Vito
San Pedro typ ef ig s wit h skin Buino, Carvalhal, Castle Kennedy, Chiajese, Dauphine , Drap
bronze or violet;pulp amber or red d 'Or(syn .RoyalVineyard),FracazzanoRosso,Lampeira,Ottato
Rosso, P etronciano , Pied de Bo euf
Comm ontype figs w ith skin A lb o, An ge lin a, B lan c he , B ri nd is ino , C as tel lan a, C itr at o,
green or yellow; pulp amber C olombo B ianco, Do rée Nobis, D ottato, E arly White, E deisi,
or white Flore n tine, Fra cazza n o , Hd a di, Hde id a rm al, Jau n e de
Toulou se, K ahraman i, Kar gigna , Mag dalen, Min uto B ianc o,
Mshak i, Ojo d e Perdiz, P alla ro, Pelo d e Buey,P oppa, Ra by
Cas tle, Re ine, Sc hifo , Seirolles , Tos sico, Urja l, Verde ccio ,
Verdillo ,Yaffawi,Zo nto,
Comm ontype figs w ith skin Abakor,Aberdin,An gélique,ArneoBianc o,Barnisso tteBlanc he,
green or yellow; pulp various Ba youd i, Bo nta rd, Bo uin, Bou krati, Br ianzo la, B utt afar ro,
shades of red Carab aset a, Ca rlin a, C ime iren ca Bla nca , C ol d e Da me,
Cotignacenq ue, Dame Blanche, Darji, Datte, Daurada, Doree,
Du R oi, El Kh adri, G enoa, G ota d e Mel, Gr assale, Gr asse,
Grosse Jaune, Ha rraki,Kaffi, Kom mathri, Limoncello, Lucano ,
Ma rseilla ise, Ma rtig iano, Meou , Mo naco , Mo nstru eus e,
Mo resca , Mus sega, Neimi, Paja jero , Pan ach ee, Per tico ne,
Pissalutto,Pounchuda,Quarai,Ravignon,Salerne,Sfari,Sicile,
Sig nor ella, Slat i, Su cra da, S ulo mo, Syda wi, T ban i, Tord o,
Toscana,Troiano,Unnequi,VerdaleBlanche,Verdeal,Verdino,
Verd one, Vert e, Vescovo , Viola da Blanca, Yellow Nech es,
Zamo zujica, Z imitza,
Contd. Table]
[Table Contd.
168Breeding of Un derutilized F ruit Crops
Comm ontype figs w ith skin Agu lla, Alican tina, Biter Abiod, Cas eta,C astagnola, Cervon e,
green or yellow; pulp color not Cistallino, Comadre, Du Japon, E spagnole, Graissane, Jo rba,
de sign ate d Levan t, Llimon enca, N ew Fren ch, Par ejal, Pelo sa, Pilosella,
Préco ce, Ragu sa, Reginella, S erra, Sextius, Su ltani,Tassiret,
Tiboulenque ,Tira , Vacal
Comm ontype figs w ith skin Abbou di,Albanera,Archipel,Barbillone,Beall,Brunswic k,Datil,
dark ( various shades o f red, Leon, Marinera, M arsaoui, Matelassa, Mwazi, Osb orn Prolific,
brown, or violet to black); pulp Regg itana, Roja , Scavello
white or amber
Comm ontype figs w ith skin A dam , A ma ro un a, A za ich , B arg em on , B ar ni ss enc a,
dark (various shades of red, B arn issot te, Bec de Per drix, Bec uelle, B ello na, B eta da,
brown, or violetto black); pulp Blavette , Bocard e, Bord eaux, Bou chebarriq ue, Bou rdissotte
various shades of red No ire, Brias ca, B riasca, Dou ssa, Brow n Tur key, Caiana,
Calabresa, C alderona , Caravanc hin a Negra, C ascitello,
Casta gnolo , Celestin e, Cern ica, Clave u, Col d e Dame Noir,
Colo mbo N ero, Con stantin e, Couc ourelle Ga votte, Cum pini,
Cuore,Cu rigo,DameNoire,DatteQuotidienne,Dattero,Djerbi,
EarlyViolet,Eva,Ferguson,FigueFleur,Ford,FranchePaillard,
Fr anc isc ana , Fr ette, G allu cc io, G hzali, G ou raud R ouge ,
Guilia na, Hma ri, Hun t, Impe rial, Ischi a Black, Ischia B rown ,
Jaspe r, Jer u sa le m, K a hili , Ku s, L ar d ar o, Le venss an a,
Made line, Malm aison, M alta, Map pafero , Marseillais e Black,
Marse illaise L ong, M artiniqu e, Meirana, Melagrano , Melinga,
Melou ba, Mer engiana, M erioun , Merlinga , Monaie, M oscate l
Pre to, Mo uisso nne, Mou renao , Napolitaine, Neg rette, No ir
Moutier,Noral,Observantine,OEildePerdrix,Paradiso,Pardo,
Pastiliere, Pea u Du re, Pec o n jud o, Pe nna, Perro quin e,
Pis salut to Nero , Po ona, Pou lette , Precoce de Bar celona,
Pregu ssata, Preston, R amsey, Rec ousse Noire, Re gina, Rei,
Roc arde, Rose B lanche, Rose N oire, Rose P eyronne, Royal,
SaintEsprit, Saint J ean, Salada, San Piero, Sarreigne, S bayi,
Selteni, Shun nari, Shu nnari Asmar, S ignora, S mari, Sof eno,
St. Do mingo, Suk kari, Su ltane, Su ltani, Tem ri, Toulousienn e,
Tributaria, Verdal, Verdald eValence, Verdal Rou nd, Vernino,
VerteBru ne, VioletSepor, Violeta Negra, Zaiti, Ziza Kheden
Source: Condit, 1955
13.1.Important FigVarietiesGrown inIndia
13.1.1.Dinkar
SelectionmadefromlocalDaulatabadtrees.Externalskincolourisdarkred;fleshcolour
isdark pinkandpear shapedfruits.Averagelengthoffruitis6.80 cm,diameter is 11.81
cm,fr uit weight is 25.76 g.Average volume is20.16 ml, specific gravityis 1.2774 m/v;
moistureis74.0%, TSSis18.36%,totalacidityis0.23%,reducingsugarsis13.40%,
Contd. Table]
Fig(Ficus carica)169
nonreducingsugarsis1.89%andtotalsugarsis15.29%.Highyieldingvarietyproducing
big sized fruits (70100 g). It is resistant to rust disease. Average yield is 6070 kg per
tree. The cultivar is recommended for Maharashtra state except Konkan region.
13.1.2.PoonaFig
Itisthemostpopularcultivargrownfor consumptionasfreshfruit. Externalskincolour
is reddish green, flesh colour is pink and pear shaped fruits, sweet and good flavor.
Averagelengthis6.58cm,Averagediameteris12.00cm,fruitweightis24.60g.Average
volumeis18.80ml,Specificgravityis1.3076m/v,Moistureis76.0%,TSS.is19.34%,
total acidity is 0.21 %,r educingsugar sis 13.76 %, non reducing sugars is 2.25 % and
total sugars is 16.01 %. Most of the fig grown in Mangalore, Bellary, Coimbatore,
Daulatabad, Ganjam, Lucknow and Sa haranpur resemblesin plant andfruit morphology
to that of Poona Fig.
13.1.3.Conadria
Firstartificialhybridfig.Externalskincolourisgreenish,fleshcolouriscreamy pinkish
and pear shaped fruits, mildly sweet. Average length is 7.07 cm, Average diameter is
13.80cm,fruitweight is38.76g.Average volumeis33.68ml,Specificgravityis1.1580
m/v,Moistureis80.0%,TSS.Is20.15%,totalacidityis0.17%,reducingsugarsis15.66
%,nonreducingsugarsis2.08%andtotalsugaris17.74%.Goodfresh,excellentdried.
MoreproductivethanAdriatic.Lightbrebacrop.Treevigorous,tendstoexcessivegrowth
under irrigation, best inhot climates.
13.1.4.Deanna
Externalskincolourisgoldenyellow, fleshcolouriscreamywhiteandbellshapedfruits.
Averagelengthis7.81cm,diameteris16.03 cm,fruitweightis46.64g,Averagevolume
is44.76ml,specificgravityis1.0478m/vandmoistureis78.0%.TSS.is21.20%,total
acidity is 0.15 %, reducing sugars is17. 25%, non reducing sugars is1.79 % and total
sugar is 19.04 %. Highest yield of dried product is 19.82 %. Dried fig could be stored
more than 180 days at low temperature maintaining itsphysicochemical characters and
high organoleptic score.
13.1.5.Excel
External skin colour is greenish yellow, flesh colour is creamy and bell shaped fruits.
Averagelengthis6.52cm,Averagediameteris11.87cm,fruitweightis24.72g,Average
volumeis21.16 ml,Specificgravityis1.1675m/v,Moistureis75.0%,TSS.Is19.36%,
total acidity is 0.19 %,r educingsugar sis 14.86 %, non reducing sugars is 1.62 % and
totalsugaris16.48%.Fruitsareverysweet,excellent,multipurposefig.Lightbrebacrop,
similar to Kadota but more productive.
170Breeding of Un derutilized F ruit Crops
13.1.6. YCD 1 T IMLA
ItisanintroductionandreleasedfromHorticulturalResearchStation,Yercaud.Treesare
well adaptedtotherainfedsituationsofShevroyshillsandtothepoorshallowandrocky
soils.Treesshowedhigh droughttolerance besidesitsexceptionallyhardynatureandfree
fromanypestordiseaseincludingthecommonfigrust.Theplantsaresphericalincanopy
and are elegant with dense dark green leavesoften growing to a height of 7.0 m with
a spread of 12.0 m. Fruits are in attractive reddish purple color and are large in size
mea suring 7.0 cm in diameter, each weighing 100200g. Higher harvests are made from
each tree, the maximum being 4000 fruits. The bearing is throughout the yea r excepting
winter months. Fruits are a rich source of vitamin C (500 mg/100g) besides â carotene
and lycopene.
13.1.7.Celeste
Pearshaped,ribbed,sometimeswith a shortneckandslender stalkto3/4in(2cm)long.
Eye(openingat apex)isclosed, fruit issmalltomedium. Skinpurplishbrown or bronze
tingedwithpurpleandcovered withbloom. Pulpwhitish orpinkishamber,ofrichflavor
andgoodquality;almostseedless.Maincropis heavybut ofshortduration.Celeste figs
are most commonly grown in Texas.
13.1.8.BrownTurkey
Broadpyriform, usually without neck, medium to large, coppercolored. Pulp is whitish
shadingtopinkorlightred,goodtoverygoodqualitywithfewseeds.Thetreeisprolific.
Themaincrop,beginninginmidJuly,islarge;theearly,breba,cropissmall.Thiscultivar
iswelladaptedtowarmclimates.Fruitsareexcellent forjams,canning, dryingor eating
fresh. It isgrown on all theislands of Hawaii.
13.1.9.Brunswick(Magnolia)
Leavesnarrowlobed;fruitsofmaincropareobliqueturbinate,mostlywithoutneck.Fruit
stalkthick,oftenswollen;fruitofmediumsize;bronzeorpurplebrown.Pulpwhitishnear
skin, shading to pink or amber, hollow in center, offa ir to good qua lity, nearly seedless.
Ripensoveralongseason.Brebacroppoor;large,bronzeskinned,fleshlightred,coarse.
13.1.10.Marseilles(WhiteMarseilles,orLemon)
Fruitsofmaincroproundtooblatewithoutneck,onslenderstalksto1/4 in(6mm)long;
of medium size. Those of breba crop, turbinate with short, thick neck andshort stalk;
yellowgreen with small green flecks; pulp white, sweet; seeds large, conspicuous.
Marseilles is commonlyfound inCalifornia.
Fig(Ficus carica)171
13.1.11.Adriatic(WhiteAdriatic, orGrosse Verte)
Turbinatewithshort, thickneckandshort stalk; abovemedium size.Green to yellowish
greenwithredpulp;ofdistinctiveflavorandverygoodquality.Inearly,minor,brebacrop
thefruits are obliquepyriform, la rge, green,often tinged with purplishredwith darkred
pulpandstrongflavor.AdriaticfigtreescansometimesbefoundinthestateofWashington.
13.1.12.Genoa(WhiteGenoa)
Pyriformorturbinate,veryfaintlyribbed;neckthickandshort,orabsentMediuminsize;
skindowny,greenishyellow;pulpgreenishwhitenearskin.Mostlyambertingedwithred;
hollow; of fair quality. Fruits of breba crop obliqueobovate with thick neck and short
stalk; yellowishgreen externally. Pulp lightred; offa ir to good quality. This is common
variety of Chile and Argentina.
13.1.13.PurpleGenca (Black Genoa;Black Spanish)
Oblong,broadatapex,narrowatbase;large;verydarkpurplewiththickbluebloom;Pulp
yellowish becoming reddish to red at the center; juicy, with sweet, rich flavor.
13.1.14.BlackIschia(BlueIschia)
An Italian heavy bearer variety; main crop is elongated pear shaped with small fruits;
Shortneckandshorttomediumstalk;large,21/2in(6.35cm)longand11/2in(3.8cm)
wide; dark purpleblack except at the apex where it is lighter and greenish; There are
many goldenflecks; skiniswhollycoatedwiththin,darkbluebloom; eyeopen,withred
violetscales;pulp is violetred,ofgoodquality.Inthebrebacrop,therearefewrlbsand
mostly indistinct; the fruit is small, about 1 1/2 in (3.8 cm) long and of the same width
at the apex; the pulp is red to greenishamber; of poor flavor. The tree is particularly
ornamental and the leaves are glossy, only shallowly 3 lobed. It is a heavy bearer.
At Saharanpur, India, ‘Brown Turkey’, ‘Bangalore’, ‘Black Ischia’ and ‘Lucknow’
are successfully grown.Around Mumbai, there is only one variety, ‘Poona’.
13.2 . Cultivars in O ther Countrie s
13.2.1.Ventura
Compact large tree, long green fruit with deep red flesh and excellent flavor. Good as
fresh or dried. Good breba crop. Late cultivar but matures well in cool areas.
13.2.2. Kadota (Ge ntile, White E ndich, Dottato)
Medium,skinisyellowishgreen,fleshamber,tingedpinkatcenter withrichflavor.Little
or no br eba crop. Tree upright, requires annual pruning to slowgrowth. Prefers hot, dry
climate for best quality. This variety is the commercial fig of California.
172Breeding of Un derutilized F ruit Crops
13.2.3. Sari Lop (
CALIMYRNA
)
A very large and delicious, smyrna type, yellow fig with amber flesh possessing a rich,
nutty flavor. Fruit is oblatespherical. The most important commercial fig grown in
California.
13.2.4.Tena
This isthe first hybrid to have ‘Calimyrna’ (Sari Lop) on both sides of its pedigree and
released in 1975. Thefruits are medium to large size with a greenish yellow skina nd a
white to light strawberry flesh. The eye is very tight and this helps prevent spoilage
during unfavorable weather. It has a verysweet, excellent flavor when fresh or dried,
best suited in hot, dry areas.
13.2.5.Alma
Thisheavybearing, verysweetfigwasreleasedfromTexasA&MUniversity’sin1975.
It has golden brown skin and very tasty, ambertan flesh. It has a small eye which
preventsspoilageduringadverseweatherconditions.Theplantisahardy,smalltreewith
a heavy main crop that ripens very late. Highly resistant tofruit rots.
13.2.6.BlackMission
Tree very large, fruits allover black purple, elongated, Flesh watermelon to pink, fairly
good taste. Easily dried at home. Breba crop is prolific.
13.2.7.SanPedro
SanPedrofigisa midseason(midFeb)variety.Thefruitsarelargeandroundandvary
from a purple to pale green skin, and has sweet, creamy pink flesh. Moderate to heavy
crops. They prefer fullsun, shelter from the wind, in a welldrained, loamy soil.
FruitcharacteristicsofthefigcultivarsBursaSivahi,Yesilguz,Bardakci,SultanSclim,
Karabakunva, Beyaz Orak and Sarilop were studied by Kaynak et al. (1998). Bursa
Siyahi had the heaviest fruits and Karabakunya the lightest. Fruit width was greatest in
Beyaz Orak, Sarilop and Bursa Siyahi, and Bursa Siyahi had the longest fruits. Bursa
Siyahi,SarilopandYesilguzwere theeasiesttopeel.Bardakci,BevazOrak,BursaSiyahi
andYesilguzhadtheleastostiolcandskincracking.Theplastidpigmentsinthefruitskin
offigarechlorophyllaandb,Bcarotene,lutein,violaxanthinandneoxanthin(Puechet
al., 1976).
Mars et al. (1998) analyzed the diversity among 22 cultivars of the southern arid
regions inTunisiausingmorphomctricstudies.Cultivar characterizationandmultivariate
analysismade on the basisof 18 physical and chemical fruitcharacters of 22 main crop
Fig(Ficus carica)173
varieties suggested that fig germplasm is diverse. It was also possible to differentiate
three cultivar groups and two distinct cultivars.
In Israel,Assaf et al. (1999) divided the best cultivars with regular yields and high
fruitquality,into4groupsonthebasisoffruitcolour:(i)greenyellowfigs(KefarUriyya,
Jafa Kadouri, Kurt, Shahmani) (Nazarati types, Nazareth and Smirna); (ii) greenviolet
figs(BigRed,Hamoudi,K 16andSbaii), (iii)brown striped figs(GzaliandHortemani)
and (iv) black figs(Ita lian Black, Haroubi (all types), Robinb andShaltouf ElAbed). It
was noted that of the spring figs, the Nazareth was best. Growing under net protection
and importing ofnew cultivars are suggested for improving quality.
Seventeenfigcultivarswereassessedintrialsat Valenzano,Italyandcv.Petrelliwas
judged best for producing first crop figs(brebas) and cv. Donato for second crop figs.
CultivarslikeFicoReginadiGioiadelColleandZingarelloNeroproduced goodcropsof
both types (Ferrara and Petruzzella, 1992). Masui Dauphine, an important Japanese
cultivar, was evaluatedfor5yearsintheZhenjiangareainChina. It producedlarge(80
100g),elongateoval,purplered fruits with15%soluble solidscontentandhigh aspartic
acidcontent(0.22mg/100g).Thefruitwassuitableforfreshconsumptionandprocessing
(Yang et al., 1994). ElKassas et al. (1992a) evaluated the growth of four local and
introducecultivarsinEgypt.ThecultivarsDiRedo,Conadria,SultaniandBlackMission
were best in terms of tree vigour.
Evaluationofyieldandqualityofsvconiumalsoshowedmarkedvariationindifferent
cultivars(ElKassasetal.,1992b).Theyieldofthebrebacrop ofDiRedo, Conadriaand
Black Mission was >15% lower than that of the second (main) crop. Other tested
cultivarsyieldednegligiblenumbersofbrebafruits.Longerbearingunits(15buds)produced
more br eba syconia. Tlic percentage of ripesyconia/shoot in Kahramani, Black Mission
and Abboudi was 83.685.2%. In Brown Turkey and Sultani this value was 76%, in
Conadria and Abiad Asswan 69% and in DiRedo 63.1%. Sultani and Black Mission
yieldedapproximately 10 kgripefruits/treeandKahramani andAbiadAsswan 35.6 kg/
tree.Most fruits ripened fromthethirdweekofJuly;thoseofSultaniandBrownTurkey
beganto maturein the second weekofAugust. Average fruit fresh weight ranged from
11.8ginAbboudito40ginDiRedo.Totalsolublesolidsandsugarcontentswerehighest
inBlackMissionandConadria(>25°/oand>17.9%respectively).BrownTurkeyhadthe
lowest values for these parameters (16 and 13% respectively).
InBertregioninAlbania,23cultivarsoffigwereevaluatedbyNici(1989).Kaliamata,
Roshnikas,Perdhikulia,AdriatikandBajuniShkodresweremostsuitableforjamproduction,
andthefirst3ofthesevarieties(togetherwithBishtkugiiZiismokthinestosomeextent)
were the best for drying. Varieties with a high sugar percentage were judged best for
makingcandiedfruitjelly,figcakeandthealcoholicspirit‘raki’.Forhighfruitproduction.
Some varietiesofthecaprifigtypeneededartificialfertilization(caprification)toproduce
a second crop.
174Breeding of Un derutilized F ruit Crops
14. FUTURE THRUST
Rich genetic diversity available in the different part of the country should be exploited
systematicallyforselectionand screeningofgenotypeswhichmaysuitablearidandsemi
arid regions too and under problematic soil. Genotype selected must have high sugar
content and better shelf life so as value added and export oriented processed products
may be developed. Diverse patterns of fruit characteristics could be the basis of a
programmeassistedby a moleculargeneticmarker approach.Moreover, thedevelopment
of varieties more adapted to industrial uses and dried fig must take in consideration.
Further studies are needed involving chemical, biochemical, and molecular markers for
estimation of genetic variation at the molecular level. In many places, threat of genetic
erosion is evident, particularly for cultivars of “Smyrna type”.Alternative methods for
genetic resources management must be considered. There should be effective strategy
forFigMosaicDiseaseasitremainsaseriouspathologicalconstraintduringfiggermplasm
exchange.
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