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LONGAN
(
Dimocarpuslongan
Lour)
D.S. Mishra, Binayak Chakraborty,
H.Rymbai,N.A.Deshkmukh,A.K.Jha,G.F.War.,D.Paul,R.S.Patel,
L.K.Mishra, D.RoyandP.Lyngdoh
1. INTRODUCTION
Longan(Dimocarpuslongan Lour.) is a
subtropicalfruitoftheSapindaceaefamily
closely related to lychee and rambutan.
It is a fruit of leading economic
importance in Thailand (Nakata et al.,
2005).Longanisanimportantarilloidfruit
tree of the rural economyin the form of
employment and income generations of
the AsiaPacific Region. The longan
resembles the litchi (Litchi chinensis).
It is thesecond most important fruit
crop interms of area andproduction in
the family Sapindaceae after litchi.
Botanical synonyms of longan include
Dimocarpus longan Lour., Euphoria
longanSteud., Euphoria longana Lam.,
and Nephelium longana Cambess. It has also been referred to as the “little brother of
the lychee” or Dragon’s eye or Eyeball (Morton, 1987). Now a day, it is becoming an
economically important crop in a number ofcountries ofAsiaPacific Region. Thisfruit
cropisalsocultivatedinQueenslandinAustraliaandFloridaandHawaiiinUSA.Similar
tolitchi,theedibleportionoflonganconstitutesfleshy,translucentwhitearilwhichencircles
a brown to black seed. In sever al countries where litchi and longans are grown, longan
has not gained so much attention as compared to litchi. However, in Thailand, longan
contributes far better than litchi in local economy (Choo, 2000).Although the fruit is
19
CONTENTS
1. Introduction...............................................255
2. Domestication ............................................256
3. Taxonomy...................................................257
4. CentersofOrigin .......................................257
5. ObjectiveofImprovement .........................259
6. Cytogenetics ...............................................259
7. InheritancePattern .................................... 259
8. ProbleminBreeding ..................................260
9. FloralBiology .............................................260
10. BotanicalDescription .................................261
11. DifferentSpecies .......................................262
12. CropImprovementMethods......................264
13. CultivarsDeveloped...................................269
14. FutureProspects ........................................ 272
256Breeding of Underutilized Fruit Crops
consideredtobeeconomicallyimportantinAsia, however,itscommercialexploitationhas
been realized only inThailand, China, Ta iwa n Province of China and Vietnam (Jiang et
al.,2002;Rangkadilok etal.,2005).ItisalsofoundgrowinginVietnam,Cambodia,Laos,
Queensland (Australia) Indonesia and Florida and Hawa ii in USA. Longangrowing has
also been extended to India, Sri Lanka and Myanmar (Anonymous, 2013).
Longan are known by various name, the word ‘longan’ or ‘long yan’ or ‘lungngan’
derivesfromtheChinese.InThailandknownaslamyai;MalaysiaandIndonesiaas leng
keng; Myanmar as kyet mouk; Cambodia as mien; Laos as lam nhai and nam nhai;
Vietnam as nhan; French as Longanier, Oeil de Dragon (Anonymous, 2013).
Longan fruits are generally consumed as fresh. The fruits can be frozen, canned or
dried. During canning, fruits are genera lly dipped in its own juice with less or no added
sugar due tohigh soluble solid content. The flavour retentioncapa city of cannedlongan
is far better than litchi and rambhutan. Dried arils of longan have smoky flavour and
usuallyused to make refreshing drink inChina. Beside drink, dried longan arils are also
used in traditional herbal medicine for stomach ache, insomnia and as an antidote of
poison.Longanliqueurispreparedbymaceratingarilsinalcohol.Thenutritionalfact in
100 gram aril of longan is given below:
Table 1. Nutritional compo sition of longa n fruit.
Nutritiona lvalue Conten t (per 100 g ) Nutritiona lvalue Conten t(per1 00g)
Calories 60 k cal P 6.07.0 mg
Moisture 72.4 83.0 % Fe 0.3.60 mg
Pro tein 1.01.3 g Vit.A 28.0 I.U.
Fat 0.10.5 g Vit. B
1 0.03 0.04 mg
CHO 15.125 .2 g Vit. B
2 0.07 0.14 mg
Fibre 0.41.1 g Niacin 0.30.6 mg
Ca 610 mg Vit.C 60.084.0 mg
(Wong and Saichol, 199 1; http://ndb.nal.usda.gov/ndb/foods/show/ Accessed Dec., 2013; Wall,
2006)
Beside fruit, longan seeds are also used as shampoo due to high saponin content.As
a traditional medicine, longan fruits are used to enhance memory (Park et al., 2010),
promotebloodmetabolism,relieveinsomnia,andpreventamnesia.Itssecondarymetabolic
products have been also shown to have antioxidative, antiobesity, anticancer, anti
tyrosinase and immune modulatory activities (Prasad et al ., 2010; Zhong et al., 2010).
2. DOMESTICATION
LonganisgrownthroughoutSouthEastAsia,arangeoftropicaland subtropical,forits
fruit(WongandSaichol,1991).Historicalevidencessuggestedthatlonganwascultivated
insouthernChinamorethan2000yearsago.InThailand,itisknownas‘lamyay’which
Longan (Di mocarp us longan Lo ur)257
was believed to be came from theChinese name ‘long yan’ or ‘lungngan’. Further, the
fruitspeciesspreadintoLaofromThailandfollowedbyVietnam.Itisapparentlycommon
inReúnion,Mauritius,Queensland(Australia)andFlorida(USA)(Lim,2013).However,
domestication of longa n from Chinato other part ofthe south eastAsia is thought tobe
veryr ecent (Blench, 2008). This fruit is considered one of the important economic fruit
crops in South Ea stAsia (Crane, 2000). Today, the growing of this crop has also been
extended to India, Sri Lanka and Myanmar (Anonymous, 2013).
3. TAXONOMY
LonganbelongstothefamilySapindaceae,subfamilySapindodeaeandthetribeNepheliaea
with six major species viz., Dimocarpus longan, Dimocarpus dentatus, Dimocarpus
gardneri, Dimocarpus foveolatus, Dimocarpus fumatus from eastern Malaysia, Sri
LankaandIndiaandDimocarpusaustralianusfromAustralia(Leenhouts,1978).Longan
(Dimocarpus longan Lour.) contains two subspecies i.e., i) ssp. Longan and ii) ssp.
Malesianus.Eachsubspecieshaveseveralvarieties.Thecommerciallongan(Dimocarpus
longan ssp. longan var. longan) belongs to subspecies longan. In Thiland, 3 different
ediblelongan types can be identifiedwithinthe varietylongan (Subhadrabandhu, 1990).
The types are:
TypeFirst:Itisalarge tree.Mostlyfoundintheforestwith smallfruitshavingvery
thin aril. It can be utilized as a breeding ma ter ia l.
Type Second: It is native to longan (“lamyai kradook” or small “lamyai”). The
branchesare almosterect.Thefruitsaresmallwithlargeseedinsideit. Mostlyfoundin
Northern part ofThailand. It is used as a rootstock for commercial longan variety.
TypeThird:Itisthecommerciallongan(“lamyaikraloke”)withlargefruitandsmall
seed.
Besidelongan,there are alsothreevarieties(viz. var.longepetiolulatus,var.obtusus
and var. magnifolius) reported to be found wild in China (Choo, 2000).
The other subspecies i.e., Dimocarpus longan ssp. malesianus contains unexploited
genetic materials having immense potential in breeding for commercial types of longan.
Thevar ieties of this subspecies can be well adoptedin unchangingheat, RH, day length
andotherclimaticconditionswhicharefrequentlyhappenedinequatorialzone.However,
recent studies indicated that Malesianus is not a subspecies of longan as revealed by
genome size analysis and trichome morphology (VanBuren et al., 2011).
4. CENTERS OF O RIGIN
Several proposed has been made on the origin of longan, which is controversial. Some
authorssuggestedthatthe originoflonganconfinedtothemountainchain fromMyanmar
throughSouthern China. However,other extends it to the north eastern, southwest India
and Sri Lanka, includingthe lowlands (Nagao and Hoa, 2000).
258Breeding of Underutilized Fruit Crops
Groff said that the longan wasintroducedinto India in 1798. However according to
the Indian literature, it revealed that this crop is native not only to China but also to
southwestern India and the forests of upperAssam and it adjoining area including the
Garo hills of Meghalaya. Furthermore, he saidthat this crop is cultivated inBengal and
elsewhere as an ornamental and shade tree (Cited in Morton, 1987).
Keetal.(2000)based onmorphologicalcharacteristicsofpollensoflongancultivars
andtheirwildspeciesinfivezonesofChinaaswellastheanalysisofbotanicalgeography
andevolution, suggested thattheprimarycentreoforiginoflongan wasYunnan,andthe
secondary centres were Guangdong, Guangxi and Hainan provinces in China.
4.1. Ce nters of Dive rsity
AlthoughthefruitisconsideredtobeeconomicallyimportantinAsia,however,itcommercial
exploitation has been realized only in Thailand, China, Taiwan Province of China and
Vietnam (Jiang et al., 2002; Rangkadilok et al., 2005). It is also found growing in in
Vietnam, Cambodia, Laos, Queensland (Australia) Indonesia and Florida and Hawaii in
USA. The longan is always given less attention in comparison to litchi. Exception in
Thailand,theeconomiccontributionoflonganexceededthatoflitchi.Longangrowinghas
also been extended to India, Sri Lanka and Myanmar (Anonymous, 2013).
Fig.1.Taxonomic hierarchyofDimocarpuslongan L our.
Longan (Di mocarp us longan Lo ur)259
5. OB JECTIVE OF IMPRO VEMENT
The preliminary emphasis in longan breeding has to be oriented towards expansion of
cultivated area. This could only be possible when a variety having a diverse climatic
adaptability.Biennialbearingisanotherproblemfacedbythelongangrowers.Smallfruit
size, less aril recovery and acid nature of juice have been placed longan after litchi in
many countries. Longan tree also encountered with some diseases and pests. Therefore,
to develop a new variety a breeder shouldconsider the following objectives:
i. Todevelopcultivarshavingdwarf treegrowing habitforeasymanagementto obtain
good fruitquality.
ii. Todevelophighyieldingandregularbearercultivars,sincelongantreesbearirregularly
and erratically making mar keting difficulties. Hence development of high yield and
regular bearing cultivars are essential.
iii. Todevelopcultivarshavinggoodfruitqualitytraitslikelargefruitsize(1820g),high
arilrecoveryandsweetjuiceinadditionto haveprolongshelflifeandpurewhitaril.
Longanhasarelativelyshortpostharvestlifewithrapid deteriorationinfruitquality
and appearance.
iv. Todevelopcultivars resistanttodiseaseslike‘witchbroom’ andblackmildew,since
noneofthelongancultivarsisknowntoresistantto‘witchbroom’disease,therefore
development of resistant variety to this malady is needed.
v. To developcultivars resistant toinsectpestslikegal mite(Eriophes dimocarpi) and
litchi stink bug (Tessaratoma papillosa).
vi. To develop cultivars with tolerantto str ongwind dama ge and strong cyclonicwinds
as these cause branchsplitting and fruit shedding.
vii. Todevelop cultiva rs having wider adapta bility to diverse ecological conditions
viii.To develop cultivars having fruit with aborted seed
6. CYTOGENETICS
Longanisadiploidwithx=15and2n=30.Liuetal.(2011)studiedthemeiosisprocess
of pollen mother cell (PMC) of‘Shixia longan’ andobserved that i) themeiosis process
of longan pollen mother cells were closely correlated with the bud size and the anther
color, and the difference inthe stages of meiosis existed in thesa me bud and anther,ii)
aseries of abnormal phenomena in meiotic division of pollen mother cells such as
chromosome fragments, lagging chromosomes, chromosome bridges, asynchronizationof
chromosomes segregation, micronucleus, deviant tetra and so on.
7. INHERITANCE PATTERN
Anextr emelyvariable exists among seedling progenies. Small fruit size appears to be a
dominant characteristic. Yang and Chen (1987) studied on the inheritance pattern of
260Breeding of Underutilized Fruit Crops
shriveled seed and found the annual variation and variation between trees of the same
cultivar. Coefficientofvariationforshriveledseedvariedfrom8.8–25.8%incultivars.
The used of embryo culture echnique has been suggested to breed large fruited varieties
with aborted seeds (Lai et al., 1997).
7.1.LinkageMap
A linkage map constructed using 94 progeny obtained from ‘Fengliduo’ x ‘Dawuyuan’,
‘Fengliduo’isahighqualitycultivar,while‘Dawuyuan’hasalargefruitsize(Guoetal.,
2010).RAPD,ISSR,sequencerelatedamplifiedpolymorphism(SRAP),andAFLPmarkers
were usedin and a pseudotestcross design. And to construct the linakge maps JoinMap
3.0wasused.Incultivar‘Dawuyuan’,251 markersweremappedinto 22linkagegroups
covering 1,064 cM. In ‘Fengliduo’, 183 loci were mapped and 21 linkage groups were
identified covering965cM.Thiswasthefirstreportedmaponlonganwhichneedstobe
enhanced with codominant markers (Arias et al., 2012).
8. PROBLEM IN BREEDING
Majorproblemsencounterinlonganbreedingareirregularfloweringandbiennialbearing,
small fruitsize, witches’ broom disease, fruit bats and largeunmanageable treesize.
9. FLORALBIOLOGY
9.1.Flowers
Inflorescence islar ge of 3050 cm length, determinate (cyme) and composed of multiple
branched leafless panicles. Three to five flowers are present in small tertiary branches
ofpaniclesi.e.,incymules. Flowerissmallwhitecupshapedbear onpaniclesfromApril
to Julya nd normally only the central flower developed into fruit. Flowers are sma ll and
yellowishbrown,calyxlobes2 5mmx 13 mm;petals5,1.5 6mmx 0.6 –2mm,
densely woolly to glabrous (Choo, 2000). Differing with litchi, longan flowers have 5
numbersofpetals.Theflowersoflonganwithinapaniclearemadeupofstaminate(pistil
non functiona l), pistillate (stamens nonfunctional) and hermaphrodite flowers (Crane et
al., 2000).Thestaminateflowerhas 8orless hairystamensarrangedin a singlerow on
alightbrown disc.Thepistillateflowerhasanthers whicharesterileand nonfunctional.
Hermaphroditeflowercontainsbicarpellatedanddenselyhairyovarywithbilobedstigma.
Normally, only one carpel develops into fruit. The stamens of the hermophrodite flower
consistof8 sessilefilamentswithanthersproducingviablepollens.Flowersare borneon
terminal inflor escence (Subhadrabandhu, 1990).
9.2.Flowering
The longan is a crosspollinated species. In order to achieve crosspollination it has
duodichogamy that is flowering in each panicle occurs in three progressive stages of
Longan (Di mocarp us longan Lo ur)261
flowersopening,whichopendirectlyaftereachother,withacertaindegreeofoverlapping.
Generally within a panicle the first phase in the sequence of opening is the staminate
flowers,followbythepistillateflowersandthenthehermaphroditeflowersandfinallythe
staminate flowers again. Male and female phases of flowering overlap 4 6 weeks
dependingoncultivars. Thereis an overlapof flowertypes onindividualtrees,sincenot
all inflorescences develop at the same time.
9.3.Pollination
There are few reports on flowering and subsequent pollination of longan (Menzel and
McConchie,1998). In Queensland, ithas beenestablishedthat pollination in longan isa
combination of autogamous (selfpollination), wind or insects (Blanche et al., 2006).
However,toobtain goodyield foracommercialcrop,pollinationbyinsects,inparticular
bythehoneybeeisconsideredcrucial(BadiyalaandGarg1990; DuToit1994;McGregor
1976;MenzelandWaite,2005)andalsoflies(syrphids,calliphorids,andmuscids)(Pham,
2012). Honeybees, Apis cerana, made up 47 – 95% to longan inflorescences (Pham,
2012).
Insect pollination occurs from early morning (8:00 a m) to midafternoon (2:00 pm),
particularlyby thehoney bee(Apis cerana).InThailand,30%increasesinlongan yields
havebeenreported as ar esult ofhoney bee pollination (DAF, 2005). Onthe other hand,
Trigona species has shown to be an effective pollinator of longan crops in Queensland
(Blanche et al., 2006). According to Blanche et al. (2006) that both honey bees and
stingless bee (Trigona spp) visited the flowers, however, the numbers of stingless bees
inflowerswerehighinnumbersascomparedtohoneybeewhichhadapositiverelationship
withfruit set. The highernumber of stingless bees might bedue to nea rbyof rainforest.
Crossedfruitaregenerally heavierandyieldsin rowswithtwocultivarsadjacenttoeach
other indica ted that atleast oneother cultivar and anabundanceof insectpollinatorsfor
pollentransferisrequiredforbetterfruitset,yieldandquality(McGregor1976).Though,
TrigonaspeciesmaybegoodpollinatorsoflonganinQueensland,butplacingofbeehives
in longan orchard leading to increase cross pollination to produce desirable yield and
quality.
9.4.Fruit
Theperiod from flowering to harvesting gener ally varies between 57 months.T hefr uits
are similar to those of the litchi, but smaller, smoother and yellowishtan in colour. It is
milder in flavor and less acidic than litchi.
10. BOTANICAL DESCRIPTION
Longan is an evergreen tree. It may grown up to a height of 20 m with dense canopy,
brittlewooda nd corky barkthat splits and peels.T he branches arespr eading orerect in
262Breeding of Underutilized Fruit Crops
nature with corky bark, depending upon genotype. Leaves are alternate and parippinate
with 69 pairs. The upper surface of leaves having dark glossy green where as lower
surface is relatively pale green in colour. Young leaf flushes are appearing in reddish
browncolourandturningtopalegreenat maturity.Terminalinflorescence(panicle)is 8
40 cmlong. Panicle emergence commences during from late December to late February
(Sritontipet al., 2013).Although,inflorescence canbeseenduringspring onnew shoots
which do not bear panicles. In a panicle as much as 80 fruits can be seen. The fruit
weight varies from 520 g. However, the commercial grade of longan rangesfrom 14
18 g. The skin of the fruit is tough and leathery which changes colour from greenish
yellowtoyellowishbrownat ripening.Tuberculesare flattenedandindistinctfromlitchi.
Thearilconstitutes6075%oftotalfruitweight.Thereisonlyone small,roundtoovoid,
reddish brown to black seed in each fruit.Aril can be easily detached from the seeds.
11. DIFFERENT SPECIES
Dimocarpus longan Lour. together with Litchi chinensis Sonn. are the most popular
memberoftheSapindaceaefamily,whichhasover2,000speciesand150genera(Menzel
and Simpson, 1991). The genus Dimocarpus is reported tocontain six major species of
trees and shrubs (Leenhouts, 1971, 1973). Out of which five speciesare foundin Asia
spreading from Sri Lanka and India to eastern Malaysia. This species are Dimocarpus
longan, Dimocarpus dentatus, Dimocarpus gardneri, Dimocarpus foveolatus, and
Dimocarpus fumatus from eastern Malaysia, Sri Lanka and India while the remaining
species,DimocarpusaustralianusisfoundtoexistsinQueensland,Australia(Leenhouts,
1978). Out of which, the most commercially cultivated species is Dimocarpus longan
where the taxon Dimocarpus longan spp. longan var. longan is commonly known as
the commercial longan (Choo, 2000).
The varietiesof Dimocarpus longan ssp. malesianusaremalesianus andechinatus.
Themalesianus varietyis nativetoSoutheastAsia(NorthernBorneo).Greatest diversity
ofthisvarietyisfoundinBorneowhere,almost3040racesareexisted(Leenhouts,1971;
VanWelzenetal.,1988).Thefruitsaresmoothtowarty.InMalaysia,fruitsofthistaxon
arecharacterized byglobosesmoothfruitwhichturnbrownwhenripe.The arilrecovery
is very low and hence, unsuitable for consumption. It is often referred as Euphoria
malasianuswhichiscommonlyknownas“matakuching”or“cat”seye.Thisformisalso
found in Borneo. One superior form having densely thick warty fruits with higher aril
recoveryisfoundinSarawak.Alltheseformscanberoughlygroupedinto 3types based
on their fruit characteristics.
11.1. Diffe rent forms on the Ba sis of Fr uit Charac ters
Isau:Fruitsareglobularandremaingreenincolourwhenripe.Treesaremainlyobserved
in river banks of Borneo valley.
Sau:It is also remain green whenripe. Fruit shape is slightlyoblong.Mostly found
in river banks of Borneo valley.
Longan (Di mocarp us longan Lo ur)263
Kakus: T his type fruits are globular and turn browna t ripening.It is mostly found
in Sarawak province in Borneo.
The variety echinatus can easily be distinguished from variety malesianus by the
presence of long spine like tubercles. The fruits of echinatus are somewhat resembles
as Rambhutan (Nephelium leppaceum).
11.2.1.Subspecieslongan
The subspecies longan is the most commonly cultivated taxon is Dimocarpus longan
ssp. longan var. longan. In Thailand, ther e are three distinct edible longan types within
the variety longan (Subhadrabandhu, 1990). First type: It is a large forest tree, small
fruits,verythinaril,possiblyofinterestforbreedingpurposes.Secondtype:Itisthenative
longan(‘lamyaikradook’orsmall‘lamyai’),growinginthenorthernpart ofThailand,an
erecttree,bearsmallfruits,largeseedsandrecommendedas a rootstockfor commercial
cultivars.Thirdtypes;Itisthecommerciallongan(‘lamyaikraloke’),producelargefruits
and small seeds.
The other three va rieties known to exist are 1) variety longepetiolulatus and 2) variety
obtusus and3) varietymagnifolius. Thethree varietiesar e found growingwild in China
(Huang,1999 cited in Choo, 2000).
11.2.2.Subspeciesmalesianus
This subspecies has a huge potential to develop into a commercial fruit variety as well
incrop improvementof commerciallongan. However,itspotentialis yetto beexploited.
The subspecies malesianus are fully adapted to the conditions of the equatorial zone of
unchangingheat,humidity,daylengthandother.Thesewerecontradictorytothecommercial
longan,adaptedonlytothesubtropicsandproducenoflowerinthetruetropics.Subspecies
malesianus consist of two varieties, viz., variety malesianus and variety echinatus.
11.2.2.1.Subspeciesmalesianusvar.malesianus
It is native to SoutheastAsia with thegreatest variationfound in Bor neo whereit might
bepossible todistinguish between 30to 40 localr aces (Leenhouts, 1971;Van Welzen et
al., 1988). The diversity of this subspecies in Sarawak in the Borneo Island has been
documentedbyWongandGan(1992)andWong(2000).Thefruitsareglobulartoslightly
oblongandsmoothtowarty.InPeninsularMalaysia,themostcommonformofthistaxon
istheonewithglobosesmoothfruits whichturnbrownwhenripe.Thisisthetrue‘mata
kuching’ or ‘cat’s eye’ which has often been identified as Euphoria malaiensis. It has
averythinarilandis hardlywortheating. ThisformalsoexistsinBorneoandSumatra.
The more superior forms are found in Sarawak, all with densely thick warty fruits and
greater aril recovery percentage. These forms can be roughly grouped into three types
264Breeding of Underutilized Fruit Crops
based on the fruit characteristics: the ‘isau’ with fruits which are globular and remain
green when ripe; the ‘sau’ with fruits which are slightly oblong and also remain green
whenripe; andthe‘kakus’withglobularfruits whichturnbrown when ripe.Theleaves,
flowersandtree formsalso differ.The‘kakus’ ismorewidespreadinSarawak,while the
‘isau’ and ‘sau’ are mainly confined to the river banks of the Rajang river and to the
Bareo valley.
11.2.2.2.Subspeciesmalesianusvarietyechinatus
Itdiffersfromthevarietymalesianusinthatthefruitshave ratherlongspinesresembling
the‘rambutan’(Nepheliumlappaceum).ThisvarietyisfoundinSabahwherethe‘kakus’
also exists (Van Welzen et al., 1988). Dimocarpus longanvar.malesianus, the mata
kuching, is a subspecies of longa n that is better adapted to tropical conditions.
Thus the true tropical longan offers the greatest opportunity for selecting superior
materialandthusoffersanattractivepossibilityoflonganbecominga newfruitcropfor
the humid lowlands throughout the tropics.
11.3.RelatedSpecies
Beside litchi (Litchi chinensis Sonn.), other related fruits under the Sapindaceae family
includetherambutan(Nepheliumlappaceum),pulasan (Nepheliummutabile), mamonilla
(Melicoccus bijugatus) and taun or dawa (Pometia pinnata).
12. CROP IMPRO VEMENT METHODS
Traditional research technologiessuch as morphology, palynology, and isozymology, etc.
havebeenimproved inlongan.Themolecularmarkerssuch as RAPD,ISSR,andAFLP,
etc. have become maintechnologies inta xonomy andgeneticr elationship studies. Single
seedling selection has been developed into combination of multiple methods such as
seedlingselection,sportsselection,andsexualcrossing,andbiotechnologicalbreeding,etc.
ThelongangermplamsarebeingmaintainedatdifferentcentresinChina,Hawaii,Florida,
Australia, Indonesia, Israel andT hailand. The Na tional GermplasmRepository of China
wasestablishedin theFujianAcademyofAgriculturalSciencesduring1981wheresofar
202 cultivars have been maintained.
Although a wide va riability is known to exist, however, no breeding work has been
reportedinIndia.Cropimprovementinlonganhasbeencarriedoutmerethroughselection
among the best existing and openpollinated seedling in China, Florida (United States),
Taiwan, Thailand, Austr alia and Vietnam. Selection is being carried out on the basis of
large fruit size, high edible portion, crisp flesh, good flavour and high sugar content. In
Thailand, longan industryis based on seedlings evolved fromcultivars introduced from
China (Watson, 1984). However, these selection cultivars are considered to be superior
over Chinese cultivars.
Longan (Di mocarp us longan Lo ur)265
12.1.Introduction
IntroductionoflongangermplasmintheUSwasinitiatedin1940sthroughtheimportation
of seeds from China. However, these seedling trees were generally unreliable in fruiting
anddifficulttopropagatebyairlayering,therefore,theyneverattainedcommercialsuccess
(CampbellandCampbell,2000).Thecultivar‘Kohala’wasintroducedfortheRareFruit
Council International, Inc. by William F. Whitmanin 1954 from Hawaii. The tree was
morereliableinitsbloomingandfruiting,lightandthinskinned, witha largefruitsize.
Inaddition,itiseasyto propagatethroughairlayering.Severaltrials andevaluationhas
beencarriedoutinThailandandSouthFloridafrom‘Kohala’,however,noselectionsever
proved to besuperiortotheoriginal‘Kohala’. Inthe regionlikeIndia,wherethegenetic
baseofthecommerciallonganindustryislimited,introductionsofsuperiorselectionsand
theselectionoflocalseedlingsmayenhancethecropimprovementprogramme(Campbell
andCampbell,2000).
12.2.Selection
Improvementof longanthroughselectionhavebeenlimiteddue to seedlingprogeniesare
extremely variable. Small fruit size found to be the dominant character and inherited
among the seedlingpr ogenies.Trees having largerfr uitsize and high a ril recovery yield
lowwithirregularityinbearing.
Aselectionprogrammewasinitiatedover30yearsagoinFujianprovince,Chinawith
an objective to select abortseeded longan cultivars. So far five promising strains with
abortseeded fruits has been identified and selected, viz., Minjiao No.1, Minjiao No.2,
MinjiaoNo.3,MinjiaoNo.4andMinjiaoNo.5(Huang etal.,2000).OutofwhichMinjiao
No.4 isthe mostpr omising due to its stable andhigher rateof seeda bortion, large fruit
size,thickandgoodqualityaril,and highyield.InGuangxiprovince ofChina,twoearly
seasonstrainsoflongan,viz.,‘EarlySeason1’and‘EarlySeason2’,werecomparedwith
‘Shixia’(DimocarpuslonganLour.cv.Shixia),anearlyseasoncultivarfortheirflowering
and fruiting characteristics. It was found that ‘Early Season 1’ and ‘Early Season 2’
bloomed two daysearlier than‘Shixia’. Fruits of ‘Early Season 1’ and ‘EarlySeason 2’
matured 14days and 10 days earlier than ‘Shixia’, respectively. Single fruit weight of
‘EarlySeason1’,‘EarlySeason2’and‘Shixia’ were11.05,10.8and9.8g, respectively.
TSSof‘EarlySeason1’and‘EarlySeason2’were19.3and19.2%,being2.7and2.8%
lower thanthatof‘Shixia’,respectively.Both strainsshowgoodpotentialas high quality
early season varieties (Pan et al., 2010).
12.3.Hybridization
AnartificialcrossinghadbeencarriedinChina,usingthreecrossingparents‘Fengliduo’,
‘Dawuyuan’and‘Shixia’suchas ‘Fengliduo’× ‘Dawuyuan’,‘Dawuyuan’בFengliduo’
and‘Dawuyuan’בShixia’ (Liuetal.,2010).ATotalof455hybridseeds wereobtained
266Breeding of Underutilized Fruit Crops
and 438 of them germinated andgrew well. Further confirmedthrough RAPD analysis
that97%oftheresultedhybridsof‘Fengliduo×Dawuyuan’weretruehybrid.Therecord
showedthattheshortestjuvenilephaseofthe hybridswasonly2.5years,and thelongest
of themwas only3. 5 years, indicatedthat the juvenile phaseof longan is not very long.
Selection for fruit quality from 143 fruiting hybrids, five outstanding individuals were
identified, viz., FD34, FD57, FD73, FD82 and FD97. Out of which FD97 was the best
asitmaturesinmidSeptemberwithhighyield,evenfruitshape,largefruitsize(average
weightis12.7g),yellowishandsemitransparentflesh,crispandtenderfleshtexture,high
SSC(23.2%)andfleshrecovery(66.7%). Thetraitsoffruitsize,SSCandfleshrecovery
of FD97 wereall better andmatured 510later than thoseof ‘Caopuzhong’, the famous
highquality and late season cultivar in Guangdong Province (Liuet al., 2010).
Longan has synchronous sexual phases duringthe flowering ofa singlepa nicle. The
first flowers to open contain anthers only (functionally male), followed by flowers with
staminodes and fertile pistils (functiona lly female), and finally flowers with a nthers and
rudimentary pistils(functionallymale)(McConchieet al.,1994).Therefore,thisassistin
control pollinationsince there is no need to emasculate the functionally female flowers
(McConchie et al., 1994).
12.3.1.Intergenerichybridization
Intergeneric hybrids have been reported to occurnatur ally inC hina between longan and
litchi,referredtoaslungly(Groff,1921citedinMcConchie etal.,1994).Sincebothlitchi
and longanhave characters of economic important that are not exist in each other. For
instant, litchi fr uit are genera lly larger and more colourful tha n longan, while longan is
resistant to erinose mite (Aceria litchi Keifer) which is a debilitating pest of litchi
(McConchie etal., 1994). The evolvingof intergener ic hybrids will reveal thelinkageof
desirable characters for further improvement of longan.
Thefirstcaseofsuccessfulintergenerichybridsbetweenlycheeandlonganwasreported
usingcommercialcultivarsinAustralia(McConchieetal.1994). Successfulfruitingand
seed setting were obtained when longan wa s the pollen plant. But, reciprocal cross (i.e.
whenlitchiservesasapollenparent)yieldsaveryfewnumberoffruits.They studied the
breedingbarrierbetweencommerciallitchiandlonganadoptingreciprocalpollination.Isozyme
analysisusingphosphoglucoseisomeraserevealedthathybridprogenyonlydevelopedwhen
litchiwasthematernalparent.Themorphologicalcharactersofthehybridsweresimilarto
the ma ternal parent except leaves were smaller. Crosses between female longan flowers
andmalelycheeflowersresultedinnofertileseeds.Someworkerbelievedthat“Lizhiben”
and“LizhiLongyan”werethehybridprogenyoflonganandlitchi.However,RAPDtechnique
disagreedtheabovestatement(TongXianget al.,1998).‘LongLi’isprobablya natural
hybridofthelonganandlitchigenus(Jia, etal.,2011).
Somatichybridization inhigher plants has come into focus since methods havebeen
establishedforprotoplastfusionanduptakeofforeignDNAandorganellesbyprotoplasts.
Longan (Di mocarp us longan Lo ur)267
Thismethodeliminatesthegeneticbarrierinremote hybridization.Novelvariantcouldbe
possible to developusing thismethod.Plantprotoplastcultureandsomatichybridization
in longanhave been paid attention first in China (Xu and Chen, 1996). Some somatic
hybridsbetweendifferentlongancultivars,‘Do’x ‘BaiDam’,‘Do’x‘BiaoKhiao’,’Biao
Khiao’ x‘Bai Dam’ have been reported in recent past (Sitthiphrom et al., 2005).
ThesecondcaseofintergenerichybridscamefromSouthChinaAgriculturalUniversity
(Zhaoetal.,2008).Female‘Shixia’and ‘Zaoshu’longanflowerswerecrossedwithmale
‘Ziniangxi’ lychee flowers. In a third cross, female ‘Ziniangxi’ lychee flowers were
crossed with male ‘Shixia’ longan flowers. Over one hundred hybr id seedlings from the
threecrosseswereobtained.TheintergenericoriginoftheF1offspringwasverifiedusing
paternal specific markers and SRAPPCR.
F
1 population of ‘Fengliduo×Da wuyuan’ was developed by crossing between a high
qualitycv.‘Fengliduo’andalargefruitsizecv.‘Dawuyuan’.Amoleculargeneticlinkage
mapwasalsoconstructedusingRAPD,ISSR,SRAPandAFLPmolecularmarkersin94
F
1 individuals selectedfrom thehybrid populationofFengliduo×Dawuyuan’(Guoet al.,
2011).Themolecularlinkagemapof‘Fengliduo’fellinto20linkagegroups,whichcontained
184markers,withanaverageintervalof6.72cMandcoveredatotaldistanceof1102.8
cM; while the molecular linka ge map of ‘Dawuyuan’ fell into 19 linkage groups, which
contained243 markers, withanaverageintervalof6.02cMandcoveredatotal distance
of 1348.3 cM
Recently, it was proved by VanBuron et al. (2011) that longli is not a Hybrid of
Longan and Lychee as Revealed by Genome Size Analysis and Trichome Morphology.
The genome sizesfor lychee (554 Mb), longan (444 Mb), ‘Malesianus’ (404 Mb), and
rambutan(339Mb)aredistinctiveandinanarrowrange.‘Malesianus’hasagenome 9%
smallerthanthatoflonganand27%smallerthanthatoflychee.Itislikelyaspeciesthat
evolved independently in northern Borneo island, and could be classified as a species,
Dimocarpusmalesianus, not a subspecies oflongan aspr esently st ated. Flow cytometry
revealed a 50% variation in genome sizes among longli varieties, with genome sizes
ranging from 450 to 678 Mb, beyond the range between longan and lychee. T hegenome
sizevar iation and distinct leaf hair morphology suggest that longliis not an intergener ic
hybrid,a nd it is likely a separate genus evolved independently. The tested cultivars with
distinctive genome sizes within D. confinis could be classified as separate species.
12.4.Biotechnology
Genetic improvement in longan through modern high technologies has achieved great
progressintermsofitscontributiontoproductionparticularlyinChina(Laietal.,2000).
During1980sisozymeswereused firsttoidentifylongancultivars(ChenandKe,1989).
Molecular markers are helpful in providing information on genetic diversity and
authenticationofcultivarsandthiscouldbeusedinfuturebreedingprogrammes.RAPDs
werethefirstPCRbasedmarkerdevelopedforlongan,helpingtoidentifygeneticdistance
268Breeding of Underutilized Fruit Crops
indifferentgroupsoflongancultivars(ChenandLiu,2001;Zhongetal.,2007 andYang
etal.,2013).GermplasmsoflonganfromTaiwan,China,andMalaysiawerecharacterized
byusingthisRAPDmarker(Yonemotoetal.,2006).Recently,highannealingtemperature
RAPD (HATRAPD) technique has been popular in characterization of longan cultivars
(Sitthiphrometal.,2005).Thisisacosteffectiveandeasymethodology thatcanbeused
toidentifyplantvarieties,aswellastoadvancetheknowledgeofbiodiversityinpreviously
uncharacterizedspecies(Wangspaetal.,2005).ApplicationofAFLPshasalsobeentried
outin longancultivars(Yietal.,2003;Ganjunetal.,2003andLinet al., 2005). ISSR
markershavealsobeendevelopedandusedforcultivaridentificationanddiversityanalysis
(Hong 2007; Chianget al., 2010 and Mariana, 2013). Transgenic plants in longa nhave
been obtainedbyusingAgrobacteriummediatedgenetic transformation technology(Zeng
et al., 2000).
Thedifferentiallyexpressedgenes betweennormaland abortiveseedsoflonganwere
identified by cDNAAFLP technology. The transcriptderived fragments (TDFs) when
compared with the database using BLAST, it was foundthat three TDFs similarity with
Ca
2+
ATPase, cellulase, and plastidic ATP/ADP transporter, respectively, whose up
regulation or downregulation expression might influence several important metabolic
pathways in developingseeds and led to seed abortion (Xi et al., 2011).
RealtimereversetranscriptasePCRisapowerfultooltoinvestigaterelevantchanges
ingeneexpressionduringplant somaticembryogenesis.Thistechnologyhasbeenapplied
duringsomaticembryogenesisinlongan(LinandLai,2010).ProfilingofmiRNAsduring
somatic embryogenesis of longan has also been constructed which may contribute
knowledge towards plant cell differentiation and zygotic embryo development in longan
(Lin and Lai, 2013). Recently, longan embryogenic callus (EC) cDNA library has been
sequencedusinganIlluminaHiSeq2000systemwhichmayprovideinformationregarding
molecular resources for longan functional genomics (Lai and Lin, 2013).
InitiallyIsozymeswere used to identifylongan cultivars in the1980s (Chen and Ke,
1989).Later,itwasfindthatthismethodwasnotveryeffectiveindistinguishingcultivars.
This lead to developing of PCRbased markers, among which RAPDs were the first
PCRbasedmarkerdevelopedforlongan,allowing theseparationofdifferentgroupsand
theidentificationofcultivars(ChenandLiu2001;Linetal.2005;Zhongetal.,2007).
ThreedifferentgroupsoflongangermplasmfromTaiwan,China,andMalaysiahavebeen
distinguished using RAPD markers (Yonemoto et al., 2006). Further, Yi et al. (2003)
distinguished11clustersamongthe46cultivarsstudiedusingAFLP.Cultivarsidentification
and diversity analysis has also been carried out with the help of ISSR markers (Hong,
2007). Recently, 384 marker s had been developeda nd verified for thisspecies using the
SSRenrichmentmethod incombinationwithpyrosequencing(Techen etal.,2010).Yang
etal.(2013)identifiedfourclonesLY21,LY47,LY48andLY52usingspecificRAPD
bands. Furthermore, they identified and characterized the difference between D. longan
and D. confinis employed SCAR ma rker s anddetected thespecific bands with expected
sizesin D. Longan but not inD. C onfinis, LY21, LY47 and LY52, respectively. This
Longan (Di mocarp us longan Lo ur)269
indicateddeveloped SCAR markers could bevery useful as a specificD.longan variety
authentication.
13. CULTIVARS DEVELOPED
There are several cultivars of longan in Southeast Asia; however, the diversity among
cultivars is lower in other regions. The following are the characteristics of the most
commonly cultiva ted varieties in the world (Choo, 2000).
13.1.Chuliang
ThiscultivarisnativetoGuangdongProvinceinChinaofsuperiorqualitylongancultivar.
Fruitsize islarge(12.016.5g).Arilconstitutesabout6974%oftotalfruitweight.The
thick,firmandfragrantarilissweetintastewithTSScontent2023 %.Itissuitable for
processing. Attractive golden yellow colour is formed in aril after drying which fetches
good marketprice.Thecultivarproduceshighandstableyield.Thecultivarwasawarded
the Gold Prize at First ChinaAgricultural Exposition during 1992 and won the title of
“FamousBrandProduce”attheThirdChinaAgriculturalExpositionduring1997andalso
at theChinaInternationalAgriculturalExposition,1999(Liuet al., 2000).
13.2.Shixia
It is another popular cultivar in Guangdong Province, China. It has a long history of
cultivationabout140years.Fruitsaresmall,butcrisp,thickandsweetarilwithTSS(19
20%) and with good flavour. It is the best cultivar for consuming as fresh fruit and
produces heavy crops regularly on unusually large panicles (Menzel et al., 1990). It is
generallyconsumedasfresh.Thesmall fruitshavesweetarilwithTSScontent1920%.
Thiscultivarissuitablefor highdensityplanting
13.3.Wuyuan
‘Wuyuan’(syn.BlackRound)isanotherimportantlongancultivarinGuangdongprovince,
China.Fruitsizeismedium (15 g)withlargeseed, softandjuicyarilofaveragequality
withTSS (1415%). Trees are high yielding. The fruit is suitable for eating fresh and
dried.Seedlingsarevigorousandconsequentlyuseful as rootstock(Menzeletal.,1990).
13.4.Fuyan
It is an important longancultivar in Fujian province, China, occupied about 90% of the
area under the province. Fruit is large sized (18 g) with thin skin, small seed and thick
crisparil.Yieldis high.Itis bestused forcanningsince thefruits havea lowTSS value
of 1516 percent (Menzel et al., 1990).
270Breeding of Underutilized Fruit Crops
13.5.Wulongling
This is another ma jor cultivar in Fujian province, China. It history of cultivation ca n be
traceduptomorethan150years.Fruitismedium insize(15g)withthickskin,goodaril
recovery and sweet flavour, TSS (2123%) (Menzel et al.,1990). It has has a distinct
alternate bea ring cropping.
13.6.Daw
‘Daw is the most popular longan cultivar of Thailand, accounted for about 73% of the
totalcultivationareaunderlonganinthecountry.Thename‘Daw’meaningearly,indicating
it earlymatur ing. Flower ing takesplace in December and fruit harvesting occurs in late
June to early July. Thus fruits fetch high premium from foreign mar kets. It is the most
regularbearerandhas noirregularbearingproblem.Inaddition,the cultivarisrelatively
free from infection by witches’ broom disease (Subhadrabandhu and Yapwattanaphun,
2000).Thearilcontentislowduetobigseed, thoughhavealargefruitsize. Thearilis
sweet andwithgood flavour,israthertoughand notas crispywhencomparedtothearil
of ‘Biew Khiew’. Fruits do not keep well on the tree and the seed may even germinate
withinthefruit.Thefruitscanbeconsumed freshorprocessed.Thiscultivarisnormally
grownintheNorthernprovinceswherethecoolwintermonthsarenecessaryforinduction
of flowering.
13.7.Chompoo
ItisanotherThaicultivar.Itisa midmaturingcultivar.Fruit ismedium size,ovalshape
withgreenishlightbrownskin.Arilisverysweetandslightpink,hencenamed‘Champoo’.
Aril content is high due to small seeds with high TSS (2122%) and pleasanta roma. It
is irregular bearer.
13.8.BiewKhiew
A latematuring cultivar of Thailand. Fruit isis round, large,brownishgreen colour and
high a rilcontent.Aril iscrispy, plea santscenteda nd sweet (TSS, 22%) andof excellent
quality.Peel isthickand,therefore,advantageoustolongershelflife.It exhibitsirregular
bearing and is susceptible to witches’ broom disorder.
13.9.Haew
It is a Thaicultivar whichis a latematuringcultivar. It flowersin late January to early
February.FruitsmatureduringmidtolateAugust.Fruitismediumtolargesize,smallseed
andaverage aril content.Arilisfirmandofgoodeatingquality. Peelis rough,thickand
therefore, advantageousfor longer shelflife. ‘Haew’ flowers easily and of high yielding.
However, it is an alternate bearer. The fruits are suitable for canning. It required cool
winter for flowering induction.
Longan (Di mocarp us longan Lo ur)271
13.10.Dang
Thefruits areharvestedduringmidJulyto earlyAugustandtherefore,classifiedinMid
maturingcultivar ofThailand. Fruitis largewith reddish brown rind.Aril content islow
duetolargeseedsize.Thefruitqualitydeclineswithmaturity.Yieldandqualityaresimilar
to ‘Daw’. ‘Dang’ is susceptible to water logging.
13.11.Baidum
Thisisalsoamidmaturingregularbearingcultivarfrom Thailand.Harvesting offruitsis
carried outduring the month of MidJulyear lyAugust. Fruit is medium size with rough
rind,smallseedandmoderatearilcontent.Arilisofacceptableflavour,crispy,verysweet
andis brightwhiteincolour.Thiscultivaris a regularbearer andcanwithstanddrought
quite well.
13.12.Talub Nak
This isa nearlymaturing cultivar whose fruits are harvested in mid tolate July. F ruitis
mediumsized,withsmallseedandhigharilcontent.Arilisbrightwhiteincolourandless
sweet .
13.13.Phetsakon
Itisanearlymaturingregularcultivaroflongan.DifferingtoothervarietiesofThailand,
itisnotrequiredcoolerclimateforinductionofflowering.Thiscultivarismainlygrowing
in central region of the countryin Samut sakhon andRatchaburi provinces.
13.14.Fengko
ItisthemostpopularcultivaroflonganinTaiwanProvinceofChina,occupiedabout98%
ofthelonganareaundercultivation.It isa good yielder. Fruitisevenly largesized.Fruit
rindisyellowishbrownwhichturnsbrighteratlowtemperature.Arilisverysweet, TSS
content20%.Fruitsattachedfirmlywiththestalk,noteasilydetachedwhichisconsidered
as a good quality.
13.15.Chingko
Another famous cultivar of longan in theTaiwan Province of China. Fruit rind is light
brownish green on ripened; however, fruit quality is easily affected by environmental
conditions.As comparedto ‘Fengko’,fruitsareeasilydetachedfromthestalkandarilis
less sweet. Therefore, it is considered to be of a poor quality fruit.
272Breeding of Underutilized Fruit Crops
13.16.Longnhan
A popular cultivar of longa n in the Mekong delta, Vietnam. It is a truly tropical longan
which can yield two crops per year.
13.17.Tieuhue
‘Tieuhue’ is also a true tropical longan which can yield three crops in two years in the
Mekong delta of Vietnam.
13.18.Longhungyen
It isa popular cultivar oflonganin theNorthern regionof Vietnam.The cultiva r thrives
only in subtropical conditions and produces only one crop per year.
13.19.Kohala
It is popular in Florida, USA. Fruits are large size, high aril content with sweet spicy
flavoured aril.P runingof panicle(2/3 of thelength)in orderto increasethe fruit size is
a common practice in this cultivar.
13.20.Egami
ThiscultivarwasoriginatedinHawaii.KonaNo.1 wasselected fromanopenpollinated
seedlingofunknownoriginatKonaResearchStation,CollegeofTropicalAgricultureand
Human Resources (CTAHR). This Kona No. 1 was named “Egami”in honour of Mr.
YosotoEgami.Thepaniclesbearfruitsinlargeclusterwithexcellentfruitsettingpercentage
(more than 50 fruits/panicle). Flower ing starts in late February and continue to the end
of March. Fruits are ripened duringAugust to ear ly October. The arilconstitutes 7075
%oftotalfruitweightwithTSScontent1822percent.Thiscultivarissuitablefordrying
(Ito et al., 2000).
BesidetheabovesaidvarietiessomevarietieslikeLongnhan,TieuhueandLonghungyen
are also popular in Vietnam.
14. FUTURE PROSPECTS
Longancanbeaprofitablecropasthereis a highdemandfor longan.Therearenumber
ofbreedinglimitationswhichneedtobetakenintoconsideration.Forinstant,breedingfor
long shelflifeas thefruitishighlyperishable,regularfloweringandfruiting,dwarf tree,
large fruitsize and early or late maturity through exploitation of the natural variability
existsin longan.
Longan (Di mocarp us longan Lo ur)273
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LONGAN (Dimocarpus longan Lour)
Longan fruits. b) peeled fruit and seed of longan .
Longan fruits. a) fruit cluster of longan .