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Cucurbit Grafting
Angela R. Davis a; Penelope Perkins-Veazie a; Yoshiteru Sakata b; Salvador López-Galarza c; Jose Vicente
Maroto c; Sang-Gyu Lee d; Yun-Chan Huh d; Zhanyong Sun e; Alfredo Miguel f; Stephen R. King g; Roni Cohen
h
; Jung-Myung Lee i
a
United States Department of Agriculture, Agricultural Research Service, South Central Agricultural Research
Laboratory, P.O. Box 159, Hwy 3 West, Lane, OK, U.S.A b National Institute of Vegetable and Tea Science,
Kusawa-360, Ano, Tsu, Japan c Departamento de Producción Vegetal, ETSIA, Universidad Politécnica de
Valencia, C<i>o</i> de Vera s/n, Valencia, Spain d National Horticultural Research Institute, Rural Development
Administration, Suwon, Rep. of Korea e AVRDC-The World Vegetable Center, P.O. Box 42, Shanhua, Tainan,
Taiwan f Instituto Valenciano de Investigaciones Agrarias, Apartado Oficial, Moncada, Valencia, Spain g
Vegetable and Fruit Improvement Center, Department of Horticultural Sciences, Texas A&M University,
College Station, TX, U.S.A h ARO, Newe Ya'ar Research Center P.O. Box 1021, Ramat Yishay, Israel i Dept.
of Horticulture, Kyung Hee University, Suwon, Rep. of Korea
Online Publication Date: 01 January 2008
To cite this Article Davis, Angela R., Perkins-Veazie, Penelope, Sakata, Yoshiteru, López-Galarza, Salvador, Maroto, Jose Vicente,
Lee, Sang-Gyu, Huh, Yun-Chan, Sun, Zhanyong, Miguel, Alfredo, King, Stephen R., Cohen, Roni and Lee, Jung-
Myung(2008)'Cucurbit Grafting',Critical Reviews in Plant Sciences,27:1,50 — 74
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Critical Reviews in Plant Sciences, 27:50–74, 2008
Copyright c Taylor & Francis Group, LLC
ISSN: 0735-2689 print / 1549-7836 online
DOI: 10.1080/07352680802053940
Cucurbit Grafting
Angela R. Davis,1 Penelope Perkins-Veazie,1 Yoshiteru Sakata,2

Salvador López-Galarza,3 Jose Vicente Maroto,3 Sang-Gyu Lee,4 Yun-Chan Huh,4
Zhanyong Sun,5 Alfredo Miguel,6 Stephen R. King,7 Roni Cohen,8
and Jung-Myung Lee9
1
United States Department of Agriculture, Agricultural Research Service, South Central Agricultural
Research Laboratory, P.O. Box 159, Hwy 3 West, Lane, OK 74555, U.S.A.
2
National Institute of Vegetable and Tea Science, Kusawa-360, Ano, Tsu, Mie 514-2392, Japan
3
Departamento de Producción Vegetal, ETSIA, Universidad Politécnica de Valencia, Co de Vera s/n
46022 Valencia, Spain
4
National Horticultural Research Institute, Rural Development Administration, Suwon 441-440,
Rep. of Korea
Downloaded By: [USDA National Agricultural Library] At: 11:18 13 August 2008
5
AVRDC-The World Vegetable Center, P.O. Box 42, Shanhua, Tainan, 74199, Taiwan
6
Instituto Valenciano de Investigaciones Agrarias, Apartado Oficial, 46113 Moncada, Valencia, Spain
7
Vegetable and Fruit Improvement Center, Department of Horticultural Sciences, Texas A&M University,
College Station, TX 77843, U.S.A.
8
ARO, Newe Ya’ar Research Center P.O. Box 1021, Ramat Yishay, 30095, Israel
9
Dept. of Horticulture, Kyung Hee University, Suwon 449-701, Rep. of Korea
Table of Contents
I. CUCURBIT GRAFTING ......................................................................................................................................51

A. Introduction ....................................................................................................................................................51
B. History ...........................................................................................................................................................52
C. Current Status .................................................................................................................................................53
D. Concerns Associated with Grafting ...................................................................................................................53
E. Economic Feasibility .......................................................................................................................................53
II. WHY GRAFT CUCURBITS? ...............................................................................................................................54

A. Disease Control ..............................................................................................................................................54
B. Cold/Heat Tolerance ........................................................................................................................................56
C. Wet/Flood/Drought/Salinity Tolerance ..............................................................................................................56
D. Increased Efficiency of Land Usage ..................................................................................................................56
E. Effect of Grafting on Flowering and Harvest ......................................................................................................56
F. Nutrient Uptake ..............................................................................................................................................57
III. PLANT PHYSIOLOGY ........................................................................................................................................57

A. Scion and Rootstock Physiology .......................................................................................................................58
B. Vigor and Shape ..............................................................................................................................................58
C. Yield and Size ................................................................................................................................................58
D. Quality ...........................................................................................................................................................59
IV. WHAT CUCURBITS ARE GRAFTED .................................................................................................................60

A. Watermelon ....................................................................................................................................................60
B. Cucumber ......................................................................................................................................................60
Address correspondence to Angela R. Davis, United States Department of Agriculture, Agricultural Research Service, South Central Agri-
cultural Research Laboratory, P.O. Box 159, Hwy 3 West, Lane, OK 74555, U.S.A. E-mail: adavis-usda@lane-ag.org.
50
CUCURBIT GRAFTING 51
C. Melons ...........................................................................................................................................................61
D. Bitter Gourd ...................................................................................................................................................61
E. Summer Squash ..............................................................................................................................................61
F. Chinese Snake Gourd ......................................................................................................................................61
G. Commonly Used Rootstocks ............................................................................................................................61
V. GRAFTING METHODS ......................................................................................................................................62

A. Tongue Approach/Approach Graft ....................................................................................................................64
B. Hole Insertion/Terminal/Top Insertion Graft ......................................................................................................64
C. One Cotyledon/Slant/Splice/Tube Graft ............................................................................................................65
D. Cleft/Side Insertion Graft .................................................................................................................................65
E. Pin Graft ........................................................................................................................................................66
F. Double Graft ..................................................................................................................................................66
G. Root Pruning ..................................................................................................................................................66
H. Automated Graft .............................................................................................................................................66
I. Acclimatization ..............................................................................................................................................67
J. Production of Rootstock and Scion ...................................................................................................................67

K. Compatibility .................................................................................................................................................67
VI. CONCLUSION .....................................................................................................................................................68
ACKNOWLEDGEMENTS ...........................................................................................................................................68
REFERENCES ............................................................................................................................................................68
option. Some seed companies now offer watermelon transplants

Due to limited availability of arable land and high market de- grafted onto squash or bottle gourd rootstocks, and some trans-
mand for off-season vegetables, cucurbits (plants in the family Cu- plant facilities offer grafting services. There have been thorough
curbitaceae) are continuously cultivated under unfavorable condi- analyses of cucurbit grafting in other countries, but the literature
tions in some countries. These conditions include environments that in English is limited. This review summarizes the state of the cucur-
are too cold, wet, or dry, or are cool low-light winter greenhouses. bit grafting industry on a global level, translating work published
Successive cropping can increase salinity, the incidence of cucurbit in many languages.
pests, and soilborne diseases like fusarium wilt caused byFusarium
spp. These conditions cause various physiological and patholog- Keywords Cucurbitaceae, watermelon, cantaloupe, cucumber, meth-
ical disorders leading to severe crop loss. Chemical pest control ods, quality, yield
is expensive, not always effective, and can harm the environment.
Grafting can overcome many of these problems. In fact, in many
parts of the world, grafting is a routine technique in continuous
cropping systems. It was first commonly used in Japan during the I. CUCURBIT GRAFTING
late 1920s by grafting watermelon [Citrullus lanatus (Thunb.) Mat-
sum. and Nakai] onto pumpkin [Cucurbita moschata Duchesne ex. A. Introduction
Poir] rootstocks. Soon after, watermelons were grafted onto bottle The phase-out of methyl bromide fumigation drives the
gourd [Lagenaria siceraria (Molina) Standl.] rootstocks. This prac- search for alternative methods of soilborne pathogen control in
tice helped control declining yield due to soilborne diseases. China
produces more than half the world’s watermelons and cucumbers vegetables. Although alternative pesticides and other physical
(Cucumis sativus L.), and approximately 20% of these are grafted. treatments are being tested and developed, grafting with resis-
Use of rootstocks can enhance plant vigor through vigorous attain- tant rootstocks offers one of the best methods to avoid soilborne
ment of soil nutrients, avoidance of soil pathogens and tolerance of diseases. Additionally, grafting can affect vegetative growth,
low soil temperatures, salinity, and wet-soil conditions. The type of flowering, fruit ripening date and quality, and provide higher
rootstock affects cucurbit plant growth, yield, and fruit quality. Cu-
curbit grafting is rare in the United States, but with continued loss yields, especially under low-temperature conditions.
of quality disease-free farmland along with the phase-out of methyl Rootstock-scion combinations reportedly affect pH, flavor,
bromide, the U.S. cucurbit industry sees grafting as an attractive sugar, color, carotenoid content, and texture of fruit. As early
52 A. R. DAVIS ET AL.
as 1949, Imazu recommended Cucurbita moschata as a root- TABLE 1

stock, since it confers resistance to fusarium wilt and improves Common and scientific names of crops used in this review and
plant vigor. He noted however, that this rootstock caused in- names of commonly used cultivars and their scientific names
ferior texture and flavor in grafted ‘Honey Dew’ (Cucumis
melo var. inodorus) fruits. Fortunately, these quality effects Common name Scientific name
are not always negative. Some rootstock-scion combinations African horned cucumber Cucumis metuliferus
increase flesh firmness, along with sugar and lycopene con- Armenian cucumber, Cucumis melo var. flexuosus
tent in watermelon (Davis and Perkins-Veazie, 2005). This snake melon
study demonstrated that quality traits can be improved by Bitter gourd, bitter melon Momordica charantia
selecting rootstock-scion combinations that complement each Bottle gourd Lagenaria siceraria
other. Burr cucumber Sicyos angulatus
It is not surprising that rootstocks have such a drastic impact Cantaloupe Cucumis melo L. var.
on the scion and scion fruit, because they can enhance plant cantaloupensis
vigor, improve disease resistance, improve tolerance of low soil Chinese snake gourd Trichosanthes kirilouii
temperatures and/or salinity, and improve attainment of soil nu- Cucumber Cucumis sativus
trients and water. Additionally, studies have shown that RNA, Figleaf gourd Cucurbita ficifolia
protein, and small molecules, some causing signal transduction, Luffa Luffa cylindrica
can translocate from the rootstock to the scion, directly affecting Melon Cucumis melo L.
scion physiology. Muskmelon, netted melon Cucumis melo var. reticulatus
Researchers in some countries have examined cucurbit graft- Oriental melon Cucumis melo L.
ing for decades, but most of this information has not been trans- Oriental pickling melon Cucumis melo var. conomon
lated into English. This review combines information on cu- Pumpkin Cucurbita maxima
curbit grafting history, methods, disease resistance, and quality Pumpkin Cucurbita moschata
issues. Reports are summarized from many countries and non- Pumpkin, summer squash Cucurbita pepo
English sources. Table 1 lists the cultivars and species mentioned Squash Cucurbita spp.
throughout this article with their common name for reference. Squash interspecific hybrid Cucurbita maxima x Cucurbita
This timely review of cucurbit grafting provides a summary of moschata
existing research as the U.S. searches for alternatives to methyl Watermelon Citrullus lanatus
bromide for soilborne pathogen control. Wax gourd Benincasa hispida
Winter melon, honeydew Cucumis melo var. inodorus
B. History Cultivar name Scientific name
Research on cucurbit grafting began in the 1920s with the ‘Bloomless’ Cucumis sativus
use of Cucurbita moschata as a rootstock for watermelon. Ini- ‘Earl’s Favorite’ Cucumis melo L. var.
tial reports by Tateishi (1927) and Sato and Takamatsu (1930) cantaloupensis
described several studies and grafting applications at Kyusyu ‘Galia’ Cucumis melo var. saccharinus
University. Tateishi (1931) reported on grafting of watermelon ‘Honey Dew’ Cucumis melo var. inodorus
onto Cucurbita moschata rootstocks, a technique that was well ‘Shin-tosa’ Cucurbita maxima x Cucurbita
known at the time. However, bottle gourd and wax gourd [Ben- moschata
incasa hispida (Thunb.) Cogn.] became preferred rootstocks for
watermelon grafting on account of their resistance to fusarium
wilt, their high affinity for watermelon, and the highly stable
growth of the plants after grafting (Sato and Takamatsu, 1930; cumis melo. L. var. conomon Makino) (Konno, 1919), makuwa
Matsumoto, 1931; Tateishi, 1931; Kijima, 1933; Murata and melon (Cucumis melo var. makuwa Makino) (Jhoya, 1938), and
Ohara, 1936; Kijima, 1938). ‘Earl’s Favorite’ (Cantalupensis Group) (Tanaka, 1942) neces-
Pumpkin and squash (Cucurbita moschata, Cucurbita max- sitated grafting of these cultivars onto resistant rootstocks. Mat-
ima Duchense ex. Lam., Cucurbita pepo L.), figleaf gourd sumoto (1931) reported that Cucurbita spp. and melon are com-
(Cucurbita ficifolia Bouché), cucumber, and bottle gourd were patible with cucumber, and Imazu (1949) noted that Cucurbita
reported as compatible with cantaloupe (Cucumis melo L. var. moschata, bottle gourd, wax gourd, and luffa [Luffa cylindrica
cantalupensis) in 1931 by Matsumoto. Compared to water- (L.) M. Roem.] were also compatible with cucumber.
melon, progress on melon grafting (Cucumis melo L.) in Japan Cucumber grafting started in Japan around 1960 to strengthen
was slow, since oriental melons (Cucumis melo L.) are rela- low-temperature tolerance and fusarium wilt resistance (Fujieda,
tively tolerant to soilborne diseases. However, the occurrence 1994). Following the advances by Ishibashi (1959; 1963), which
and spread of fusarium wilt in pickling melon ‘Etsu-uri’ (Cu- led to improved grafting techniques and the incubation of grafted
seedlings, grafting cucumber became common in Japan in the Spain grows around 30 million grafted watermelon plants,
1960s (Sakamoto, 1965). approximately 50% of their crop (Hoyos, 2004). Twenty mil-
Around the same time, grafting was explored in Europe. In lion watermelon and 5 to 6 million melon plants are grafted in
France, research on cucurbit grafting started in the 1950s, with Italy (Amadio, 2004) annually.
the grafting of cucumber and melon scion onto figleaf gourd Grafting is considered a good alternative to methyl bromide
to control fusarium wilt. During the 1960s melon plants were for controlling cucumber soilborne diseases in Greece (Pavlou
grafted onto Benincasa spp., to offset the effects of low soil et al., 2002). The United Nations Industrial Development Orga-
temperatures in early greenhouse grown melons (Alabouvette nization (UNIDO) recently equipped Honduras and Guatemala
et al., 1974; Louvet, 1974; CTIFL, 1985). In Spain, research on with 60,000 m2 of specialized greenhouses to produce 30 million
grafted melon and watermelon started in 1976 and was com- grafted watermelon and melon plants. These greenhouses are ca-
mercialized there by the end of the 1980s, concomitant with the pable of supplying 30% of the melon transplants in both coun-
spread of commercial nurseries (Miguel, 1986; Garcı́a-Jiménez tries (Amadio, 2004). It is estimated that approximately 2,000
et al., 1990; Miguel, 1993). Cucumber grafting in Spain be- ha of watermelon and 500 ha of melon are grown with grafted
gan in the mid-1990s, but is increasing in importance (Hoyos, plants in Honduras, and 1,000 ha of grafted watermelon and 50
2001). ha of grafted melon are grown in Guatemala (Camacho, 2007).
At the end of the 1980s melon and watermelon grafting exper- Morocco, Mexico, Cuba, and the U.S. have recently started look-
iments were performed in Italy (Trentini and Maiolli, 1989). This ing at grafting as a viable option for cucurbit crops (Besri, 2004;
technology was again introduced during the 1996–97 season as Pérez-Montesbravo, 2004), but at least 250 ha of grafted wa-
a methyl bromide alternative (Leoni et al., 2004). termelon were planted in 2007 for the U.S. market (Chadwell,
2007) and approximately 1,000 ha of grafted watermelon and
C. Current Status 100 ha of grafted melon are grown in Mexico (Camacho, 2007).
In 1998, approximately 95% of watermelon and oriental mel- Ma (2008) estimated that about 10–20% of China’s watermelons
ons were grafted onto squash or gourd rootstocks in Japan, Ko- are grafted (about 2-3M ha.).
rea, and Taiwan (Lee and Oda, 2003). Table 2 demonstrates the
latest statistics on cucurbit grafting volume and the percent of D. Concerns Associated with Grafting
cucurbits grafted in multiple countries. France and Spain cur- The main problems associated with grafting are the time
rently raise approximately 1% to 3% of their melon and cu- and labor required, cost, rootstocks rendered ineffective by oc-
cumber crops using grafted plants (Erard, 2004; Hoyos, 2004). currence of newly migrated soilborne diseases or pests, and
changes in fruit quality. Grafted seedlings are more expensive
TABLE 2 than nongrafted seeds and seedlings. It is difficult for farmers to
The most recent published data for amount of area under provide the intensive care required to raise newly grafted plants,
cultivation of select cucurbit crops and percentages of total often requiring the added cost of a transplant facility that has
crop grafted within stated country healing chambers and trained personnel.
Changes in resistance to diseases or pests must be taken into
Area under Grafted seedlings
account as well as changes in fruit quality, which occur with
cultivation used (%)
some rootstocks. These problems should be regarded as impor-
Country Crop (ha) Estimated values
tant research topics, given that grafting cultivation is an indis-
South Korea Watermelon 23,179 98 pensable production technique for sustainable agriculture. They
South Korea Cucumber 5,853 95 are mentioned in detail below.
South Korea Oriental melon 7,077 95
France Melons not reported 1–3 E. Economic Feasibility
France Cucumber not reported 1–3
The labor required, and the high cost of grafted seedlings
Spain Melons not reported 1–3
make cucurbit grafting expensive. It is difficult for farmers
Spain Cucumber not reported 1–3
who wish to perform the grafting to maintain adequate hu-
Honduras Watermelon 2,000 not reported
midity and low light levels while grafted plants heal. How-
Honduras Melons 500 not reported
ever, since some transplant facilities now offer grafting services
Guatemala Watermelon 1,000 not reported
and with improved grafting techniques and mechanization, pur-
Guatemala Melons 50 not reported
chase of grafted seedlings is becoming more affordable. Grafted
United States Watermelon 250 not reported
seedlings are more expensive than non-grafted seedlings, re-
Mexico Watermelon 1,000 not reported
flecting the cost of the rootstock seeds, which are often F1 hy-
Mexico Melons 100 not reported
brids or triploids. There are added costs involved in the grafting
Source: Erard, 2004; Hoyos, 2004; Camacho, 2007; Chadwell, 2007, operation: raising the grafted seedlings, and transporting them
Huh, 2007. to growers’ fields (Taylor et al., 2006). Additionally, grafting
decreased the transplant survival rate by up to 60% in high- and there are additional claims of improved tolerance to some
wind conditions. The resultant loss of transplants and the need foliar fungal and viral pathogens.
for replanting greatly increases the cost of grafting in high-wind Fusarium wilt, caused by Fusarium oxysporum Schltdl., is
areas (Davis and King, 2005; King and Davis, 2006). probably the most common and damaging soilborne disease
In Japan, grafted seedlings are almost four times the cost of cucurbit crops worldwide. Grafting for fusarium resistance
of seeds (0.4 US dollars per seed vs. 1.6 U.S. dollars per started in Japan in the 1920s to control F. oxysporum f. sp.
grafted seedling) (Sakata, 2008). In the U.S. grafted seedless niveum Snyder & Hansen. The first rootstock used for water-
watermelon transplants cost $1,743 more per hectare than non- melon were Cucurbita moschata, but bottle gourd soon became
grafted seedless plants (Taylor et al., 2006). That is a differ- the preferred rootstock (Tateishi, 1927; Sato and Takamatsu,
ence of around $0.50, or three times more per grafted plant. In 1930; Kijima, 1933; Murata and Ohara, 1936; Sakata et al.,
Spain, grafting costs are 0.25–0.35 Euros per plant, not including 2007). Possibly because of the widespread use of bottle gourd
seed cost. This cost is similar in other Mediterranean countries rootstocks, there were reports of Fusarium spp. isolated from
(López-Galarza, 2008). infected roots (Sato and Ito, 1962), which was later identified as
F. oxysporum f. sp. lagenariae Matsuo & Yamamoto (Sakata
II. WHY GRAFT CUCURBITS? et al., 2007). Selection for fusarium wilt resistance in bottle
gourd rootstocks, and the adoption of squash interspecific hy-
The primary motive for grafting cucurbits is to avoid damage
brids (Cucurbita maxima × Cucurbita moschata) that demon-
caused by soilborne pests and pathogens when genetic or chem-

strate immunity to fusarium wilt, has prevented the reoccurrence
ical approaches for disease management are not available (Oda,
of this resistance breakdown. Additional rootstocks were identi-
2002a;b). Grafting a susceptible scion onto a resistant rootstock
fied that confer high levels of fusarium resistance, including Cit-
can provide a resistant cultivar without the prolonged screening
rullus spp. (Huh et al., 2002) and several different Cucumis spp.
and selection required to breed resistance into a cultivar. Further-
and Cucurbita spp. (Igarashi et al., 1987; Trionfetti-Nisini et al.,
more, grafting allows rapid response to new pathogen races, and,
1999; Hirai et al., 2002). Use of these rootstocks has controlled
in the short-term, provides a less expensive and more flexible
fusarium wilt in cucumber (Komada and Ezuka, 1974; Pavlou,
solution for controlling soilborne diseases than breeding new re-
2002; Tjamos et al., 2002), melon (Imazu, 1949; Bletsos, 2005;
sistant cultivars. In addition, grafting may enhance tolerance to
Xu et al., 2005c) and bitter gourd (Momordica charantia L.)
abiotic stresses, increase yield, and result in more efficient water
(Lin et al., 1998), in addition to watermelon. Grafting to control
and nutrient use; extend harvest periods, and improve fruit yield
fusarium wilt in watermelon and cucumber is a common prac-
and quality (Shimada and Moritani, 1977; Romero et al., 1997;
tice, but it is less common in melon. Control of F. oxysporum
Oda, 2002a;b; Trionfetti-Nisini et al., 2002; Lee and Oda, 2003;
f.sp. melonis Snyder & Hansen in melon has traditionally been
Rivero et al., 2003, Hang et al., 2005). Rootstocks can also
accomplished using host plant resistance (or “resistant culti-
influence temperature tolerance, moisture extremes (drought,
vars”) making grafting unnecessary (NIVOT, 2001; Yoshioka,
flooding), and salt stress (Matsubara, 1989).
2001). However, with the increased prevalence of race 1,2, graft-
A. Disease Control ing gives an economic advantage wherever race 1,2 is endemic
In the U.S., crop rotation has been effective for cucurbit pest (Hirai et al., 2002; Trionfetti-Nisini et al., 2002).
control for many years. However, many soilborne pathogens can A recent report showed that grafting melon onto squash inter-
survive in the soil for up to 10 or more years without a host, lim- specific hybrids can provide resistance not only to F. oxysporum
iting the effectiveness of rotation for some diseases. Fumigation f. sp. melonis race 1,2, but also to Didymella bryoniae (Fuckel)
is effective at controlling persistent soilborne diseases, but the Rehm, the causal agent of gummy stem blight (Crino et al.,
phase-out of methyl bromide as a soil fumigant has made the 2007). It was pointed out that while gummy stem blight is typi-
search for alternatives necessary. In intensive farming regions, cally considered a foliar disease, D. bryoniae is a soil-inhabiting
such as Asia and Europe, managing the potential buildup of soil fungus, and one of the primary sources of infection is through
pathogens is a primary concern. Europe adopted methyl bro- the crown. It was speculated that providing resistant rootstock
mide to control soilborne pathogens; while many parts of Asia on susceptible scions prevents primary sources of infection, re-
used grafting for this purpose. Since the phase-out of this fu- sulting in reduced disease incidence.
migant, grafting emerged as an effective alternative for cucurbit Monosporascus vine decline, caused by the soilborne
crop production in Europe. In the U.S., however, grafting has not pathogen Monosporascus cannonballus Pollack & Uecker, is
emerged as a viable alternative. One reason may be that methyl a serious melon disease that, in some regions, surpasses fusar-
bromide continues to receive critical use exemptions for cucurbit ium wilt in pervasiveness (Lobo, 1990; Buzi et al., 2002; Fer-
and solanceous crops (King et al., 2008). Most grafting litera- rer, 2003). It can also cause significant watermelon disease as
ture on soilborne diseases comes from studies conducted in the well (Garcı́a-Jiménez et al., 1994; Gennari et al., 1999; Buzi
Mediterranean region and Southeast Asia. It has been reported et al., 2004). Host plant resistance to monosporascus vine de-
that grafting can improve resistance or tolerance to more than ten cline has yet to emerge as a viable option, and with reduced
different fungal, bacterial, and nematode soilborne pathogens, chemicals available, grafting is part of an integrated approach
to controlling this disease. Disease incidence on grafted plants rotiodes isolates have been identified that infect watermelon and
is not as predictable for monosporascus vine decline compared commonly used watermelon rootstocks (Shishido et al., 2006),
to fusarium wilt resistance; however, grafting alone will reduce black root rot may become a problem on watermelon and pump-
disease severity depending on the rootstock / scion combina- kin in the future. Summer cucumber production was complicated
tion, time of year, and primary inoculum levels (Edelstein et al., by severe damage by P. sclerotiodes when Cucurbita moschata
1999; Cohen et al., 2005). The reason for these variable re- rootstocks were used in Northern regions of Japan (Shishido
sults appears to be lack of true resistance in melon rootstocks et al., 2006).
to M. cannonballus. These rootstocks are instead tolerant of ‘Bloomless’ rootstocks alter several characteristics of grafted
the pathogen; their large root systems and the vigor afforded cucumbers, including their susceptibility to plant diseases. These
the scion allows for crop development in the presence of this rootstocks caused increased powdery mildew damage (Po-
pathogen (Cohen et al., 2000). Grafting along with additional dosphaera xanthii [Castagne] U. Braun & N. Shishkoff) and
control strategies to reduce pathogen populations appears to be Corynespora leaf spot (Corynespora cassicola (Berk. & M. A.
a key factor in reducing the disease severity of monosporascus Curtis) C. T. Wei) on cucumber scions (Hasama, 1992; Hasama
vine decline. Watermelon tolerance to this disease has improved et al., 1993).
through grafting onto tolerant rootstocks (Peris, 2004). Root knot nematodes can cause serious disease in cucurbit
Phytophthora blight, caused by Phytophthora capsici Leo- crops. Grafting reduced nematode gall formation in cucumber
nian, is a serious threat to cucurbit production worldwide. Cu- (Giannakou and Karpouzas, 2003), watermelon (Miguel et al.,
cumber, melon, watermelon, pumpkin, and squash are suscep- 2005), and melon fields (Siguenza et al., 2005). In some cases,
tible, and the host range includes 49 different plant species the rootstocks appear to provide tolerance by providing extensive
(Babadoost, 2005). Identification of genetic resistance to this root area and vigor (Giannakou and Karpouzas, 2003; Miguel
disease has proven difficult, but certain rootstocks appear to be et al., 2005), but some rootstocks have genetic resistance that
more tolerant than others. Takahashi and Kawagoe (1971) found is exhibited in the grafted plants (Hagitani and Toki, 1978;
that certain rootstocks reduced phytophthora blight on cucum- Siguenza et al., 2005; Gu et al., 2006).
bers, and Wang et al. (2004) showed improved cucumber toler- Increased tolerance to viral complexes (CMV, WMV-II,
ance to this disease through grafting. It is probable that improved PRSV, and ZYMV) of grafted seedless watermelons compared
tolerance to phytophthora blight is all grafting can achieve, since to nongrafted controls was reported by Wang et al. (2002). An-
no cucurbit compatible rootstock with genetic resistance for P. other report demonstrated that grafted melon plants had im-
capsici has been reported (Babadoost, 2005). The best control proved tolerance to Melon Necrotic Spot Virus (MNSV) (Wang
option at present is to combine grafting with tolerant rootstocks et al., 2002). Similarly, in grafted tomato, plants were more tol-
with cultural practices to mitigate disease pressure. erant to Tomato Yellow Leaf Curl Virus (Rivero et al., 2003).
Verticillium dahlia Kleb. is another common soilborne It is presumed that the increased tolerance is due to increases
pathogen that affects cucurbit crops in many regions. Grafting in vigor, photosynthesis, chlorophyll content, and/or peroxidase
for control of verticillium wilt was studied for cucumber, melon, activity associated with the grafted plants.
and watermelon (Alabouvette et al., 1974; Paplomatas et al., Two reports claim increased viral susceptibility of grafted
2002). While immunity to V . dahlia was not found in any root- cucurbits compared to nongrafted plants. Iwasaki et al. (1996)
stocks, some were tolerate, and grafting onto these rootstocks reported grafted cucumbers to be more susceptible to certain
can delay expression of disease for up to 20 days (Paplomatas combinations of viral infection, such as Cucumber Mosaic
et al., 2002). This delay, when combined with additional control Virus (CMV), Watermelon Mosaic Virus II (WMV-II), Papaya
strategies, will likely result in effective control of verticillium Ringspot Virus (PRSV), and Zucchini Yellows Mosaic Virus
wilt. While few reports demonstrate the effectiveness of combin- (ZYMV). These specific viral combinations caused severe wilt-
ing grafting with other control methods in cucurbit crops, high ing in grafted plants when compared to nongrafted controls.
levels of control have been achieved in eggplant when grafting Boubourakas et al. (2004) reported one instance in which grafted
was combined with soil treatments designed to lower inoculum watermelon was infected by Cucumber Green Mottle Mosaic
pressure (Ioannou, 2001). Virus (CGMMV), while nongrafted plants were asymptomatic.
Phomopsis sclerotiodes Kesteren commonly causes black Possible mechanisms explaining an increase in susceptibility to
root rot in greenhouse-grown cucumbers (Cappelli et al., 2004), viruses have not been investigated. It is possible that a lack of
and has also been shown to cause disease in melon, pump- graft compatibility weakens the scion, making grafted plants
kin, bottle gourd and watermelon (Shishido et al., 2006). The more susceptible to viral infection, or plants could be infected
pathogen can survive in soil and soil-less media and even on during the grafting operation. It will be interesting to determine
plastic containers, making control difficult. Grafting was shown if new virus-resistant rootstocks, such as Ling and Levi’s (2007)
to be an effective control measure for greenhouse grown cucum- ZYMV-Florida strain-resistant bottle gourd rootstocks will con-
bers, even surpassing chemical control treatments in some in- fer virus resistance to the scion.
stances (Wiggell and Simpson, 1969), as long as highly resistant Disease-resistance mechanisms afforded by grafting are not
rootstocks were used (Alabouvette et al., 1974). Since P. scle- well understood. It is considered that fusarium wilt resistance
is due to the inherent resistance of the rootstock, whereby the tolerance is related to higher antioxidative ability and membrane
roots exclude the pathogen and prevent infection. However, Lee stability. Grafted watermelon seedlings under low temperature
(1989) observed that adventitious rooting of the scion did not stress have higher antioxidants and antioxidative enzyme activ-
always increase susceptibility. This observation was supported ities in leaves than self-rooted watermelon seedlings (Liu et al.,
by a report stating that substances associated with fusarium tol- 2003a; Liu et al., 2004a).
erance are synthesized in the roots and translocated to the scion
through the xylem (Biles et al., 1989). However, the author’s C. Wet/Flood/Drought/Salinity Tolerance
observations revealed that in most instances, adventitious root- Grafting bitter melon onto luffa rootstock improved scion
ing of the susceptible scion will result in symptom development flood tolerance (Liao and Lin, 1996). Nongrafted control water-
in fusarium-infested soils. In the case of monosporascus vine melon and watermelon grafted onto wax gourd showed greater
decline, phytophthora blight, and verticillium wilt, the root- resistance to drought stress than watermelon grafted onto bot-
stocks were not resistant but were tolerant to the pathogens tle gourd (Sakata et al., 2007). A Cucurbita moschata cultivar,
involved. In these instances it is presumed that the large root ‘Higata 2’, performed well even in waterlogged fields (Toki,
area and increased vigor of the rootstock allow scion devel- 1972).
opment in the presence of the disease. This may also explain Rootstocks can decrease the accumulation of Cl− and Na+ in
why some researchers report an increased tolerance to certain Cucumis melo scion leaves (Romero et al., 1997). This may
foliar diseases. Understanding the mechanisms of disease re- be due to the exclusion or decreased absorption of Cl− by
sistance is important for their best utilization. When grafting is the roots and replacement or substitution of total K+ by to-
used to improve tolerance to diseases such as monosporascus tal Na+ in the leaves. Yang et al. (2006) demonstrated that
root rot or verticillium wilt, additional control strategies are rec- grafted cucumber plants have higher net photosynthesis, stom-
ommended to decrease the pathogens levels to a manageable atal conductance, and intercellular CO2 concentrations under
number. NaCl stress than self-rooted plants. Under salt stress, water-
melon grafted onto salt-tolerant rootstock demonstrated im-
B. Cold/Heat Tolerance
proved vigor and yield, and maintained good fruit quality com-
One of the most striking effects of grafting is the in- pared to watermelon-watermelon-grafted controls (Liu et al.,
creased absorption of ions by grafted plants at low temperatures 2003b; 2004b;c). It was suggested that increased salt tolerance
(Tachibana, 1982; Ahn et al., 1999). This enhanced mineral up- of grafted watermelon is linked to increased peroxidase activity
take appears to be closely associated with the activity of enzymes and decreased superoxide dismutase activity (Liu et al., 2003a;
responsible for absorption (Ahn et al., 1999). Ahn further sug- 2004a,b).
gested that increased water absorption capacity of figleaf gourd
root systems conferred cold root temperature protection to cu-
cumber scions. D. Increased Efficiency of Land Usage
Grafting is used to advance the planting date for water- Grafting can overcome problems of continuous cropping in
melon during cool periods (Taussig et al., 1996; Miguel et al., protected culture, increasing land use efficiency. In continuous
2004). ‘Shin-tosa’-type (an interspecific hybrid, Cucurbita max- cropping systems, soilborne diseases accumulate over time (Lu,
ima × Cucurbita moschata) rootstocks show superior growth 2002; Zhang and Ge, 2002; Wang et al., 2004; Xu et al., 2004;
under low-temperature conditions. Cucurbita moschata are not Feng et al., 2006b; Gu et al., 2006; Zhang et al., 2006a). In ad-
as resistant as ‘Shin-tosa’ (Okimura et al., 1986), but are more dition, grafted plants with strong root systems can adapt to less
resistant than wax gourd and watermelon. Cucumber plants optimum soil conditions such as increased salt or low tempera-
grafted onto figleaf gourd are more cold-tolerant than cucumber- ture, and have a wider harvesting period (Zhang, 2002; Liu et al.,
‘Shin-tosa’-grafted plants (Fujii, 1970). Marukawa and Takatsu 2004b; Hu et al., 2006; Yang et al., 2006). The productivity of
(1969a) examined 81 Cucurbita spp. accessions for use as root- grafted cucumber in solar greenhouses has been extended from
stocks under low-temperature conditions (minimum tempera- 3–4 months to 7–8 months during winter months (Hang et al.,
ture 8–10◦ C). They found that figleaf gourd resulted in the earli- 2005). In Northern China, nearly 100% of cucumbers and wa-
est harvest and greatest yield. Under low temperature conditions, termelons grown in solar greenhouses are grafted (Hang et al.,
grafted cucumber have higher net photosynthesis, stomatal con- 2005).
ductance, and intercellular CO2 concentrations than self-rooted
plants (Zhang, 2002; Hu et al., 2006). E. Effect of Grafting on Flowering and Harvest
Early reports on cold tolerance mechanisms, state that grafted Enhanced flowering was noticed in wild cucurbit species
cucumber plants with increased total lipids per gram fresh grafted onto squash interspecific rootstock (Nienhuis and Lower,
weight, high unsaturated fatty acids, an increase in fatty acid to 1979; Nienhuis and Rhodes, 1977). Watermelon grafted onto
total lipid ratio, and a reduced ratio of esterol to total lipids, had bottle gourd caused early formation of female flowers when
higher low temperature tolerance (Horvath et al., 1983; Bulder compared to other rootstocks (Kurata, 1976b; Sakata et al.,
et al., 1990; 1991a;b). More recently it was reported that chilling 2007). In contrast, Yamasaki et al. (1994) reported that flowering
is delayed in watermelon grafted onto bottle gourd rootstocks. mation efficiency, CO2 conductivity, dark reaction activity, and
The differences in flower initiation are negligible when suitable photosynthetic rate in the scion (Sun et al., 2002; Zen et al.,
temperatures occur at the beginning of the growing season, but 2004; Qi et al., 2006; Wei et al., 2006;). Increased photosyn-
with less than optimal conditions, grafting to these rootstocks thesis can often be realized under less than optimal growing
can delay flowering for up to one week, resulting in an equal conditions, such as weak sunlight and low CO2 content in so-
delay in fruit maturity (Miguel, 1997; Xu et al., 2005d). lar greenhouses during winter months, allowing grafted plants
Blooming is caused mainly by emission of SiO2 from tri- to produce higher yields and sometimes improved fruit quality
chomes, which appear on the fruit epidermis and cover the (Xu et al., 2005d; Hu et al., 2006; Xu et al., 2006a; Zhu et al.,
pericarp (Matsumoto, 1980; Aoyagi et al., 1986; Yamamoto 2006).
et al., 1989). Cucumber varieties grafted onto a squash inter-
specific hybrid rootstock inhibited flowering (Satoh, 1996). It III. PLANT PHYSIOLOGY
was suggested that the root may control floral transition by
Cytokinins, a group of plant hormones principally synthe-
the production of inhibitory factors in some day-neutral Cu-
sized in roots, significantly influence cucurbit plant growth.
curbitaceae plants. Blooming of cucumber fruit is almost com-
Plants with vigorous root systems release more cytokinins into
pletely suppressed when seedlings are grafted onto the cultivar
the ascending xylem sap, resulting in increased yield (Kato and
‘Kitora’ (Cucurbita moschata) (Sakata et al., 2008). ‘Bloom-
Lou, 1989). For example, cucumber plants grafted on pump-
less’ fruits have a distinct appearance and a longer shelf life,

kin rootstocks demonstrated a 2.2 times higher trans-zeatin root
features that are in demand, thus allowing their sale at a higher
content, and were higher in dry matter than self-rooted cucumber
price. This caused a rapid shift from conventional rootstock cul-
(Seong et al., 2003).
tivars to rootstocks for ‘Bloomless’ production (Sakata et al.,
Cytokinin composition varies greatly between species. For
2007).
example, only zeatin and dihydrozeatin are found in cucumber,
whereas large quantities of isopentenyladenine and isopentenyl
F. Nutrient Uptake adenosine are found in squash and gourd (Table 3). However,
One advantage of grafting is the utilization of the root- the stem must play a role in cytokine production or transport
stock’s vigorous root system. Grafting influences absorption and since a small portion of scion stem, approximately 15 to 20 cm
translocation of phosphorus, nitrogen, magnesium, iron, and cal- in length, can effectively modify the composition of cytokinins
cium (Gluscenko and Drobkov, 1952; Masuda and Gomi, 1984; in the ascending xylem sap (Lee et al., 2001a;b).
Ikeda et al., 1986; Kim and Lee, 1989; Ruiz et al., 1997; Nie Messenger RNA (Ruiz-Medrano et al., 1999; Xoconostle-
and Chen, 2000; Pulgar et al., 2000; Zhang, 2002; Hu et al., Cázares et al., 1999) and protein (Golecki et al., 1998; Gómez
2006; Yang et al., 2006). Absorption and translocation of other et al., 2005) from a rootstock can migrate through the phloem
micronutrients such as iron and boron are also influenced by to the scion to accumulate in the phloem and apical tissues.
the rootstock (Brown et al., 1971; Gomi and Masuda, 1981; It was suggested that RNA may move within the phloem as a
Zaiter et al., 1987; Rivero et al., 2004). Concentrations of N, component of a plant information superhighway (Xoconostle-
P, Ca, and Mg in exudates increased in grafted plants (Nie and Cázares et al., 1999). One mRNA, which was demonstrated to
Chen, 2000). Ion influx allows increased light energy transfor- circulate through the phloem from a pumpkin rootstock to a
TABLE 3
Cytokinin composition in xylem sap collected from intact and grafted plants of cucumber, squash, and figleaf gourd plants
Cytokinin content (ng/ml sap)
Crop Zeatin Dihydo-zeatin Isopentenyl

(Scion/rootstock) Zeatin riboside riboside adenine Total
Cucumber (Cucumis sativus) 0.076 4.55 0.80 Trace 6.11
Squash A (Cucurbita moschata) Trace 3.67 0.43 3.63 7.73
Squash B (Cucurbita maxima) Trace 4.06 0.57 1.84 6.47
Figleaf gourd (Cucurbita ficifolia) Trace 4.54 1.48 6.18 12.2
Cucumber/Cucumber 0.55 5.58 0.96 Trace 7.07
Cucumber/Squash A 1.65 4.29 0.20 Trace 6.14
Cucumber/Squash B Trace 5.36 0.19 Trace 5.55
Cucumber/Figleaf 1.49 5.08 0.65 Trace 7.22
Reprinted with permission from Horticultural Reviews (Lee and Oda, 2003).
cucumber scion, is a member of a RNA family shown to af- B. Vigor and Shape
fect apical meristem development (Ruiz-Medrano et al., 1999). Most studies indicate that changes in the scion are con-
Tiedemann et al. (1994) studied the phloem proteins that differed trolled by the rootstock through controlled uptake, synthesis,
in Cucumis sativus grafted on Cucurbita ficifolia or on Cucurbita and translocation of water, minerals, and plant hormones (Lee,
maxima. They found four proteins on SDS-PAGE gels in grafted 1994). Yetisir and Sari (2004) reported that out of 11 root-
scions that do not appear in control plants, but matched the rootstocks tested on watermelon, all the grafted plants produced
stock phloem protein pattern. Two of the proteins matched the more biomass than control plants.
phloem protein 1 and phloem protein 2 molecular weights of the The scion variety obviously affects final size, yield, and qual-
rootstock ity, but rootstock effects may drastically alter these characteris-
tics. Tamada (1989) and Hagihara (2004) classified the species
and cultivars of the rootstocks used in Japan on the basis of
A. Scion and Rootstock Physiology plant vigor. In general, ‘Shin-tosa’ rootstocks are more vigorous
The sequence of structural events involved in graft compat- than the bottle gourd rootstock cultivars. Most of the Cucurbita
ibility of herbaceous plants begins with ruptured cells at the moschata cultivars are comparatively weak. Cultivars of Cu-
graft interface collapsing to form a necrotic layer that disappears curbita pepo and watermelon range from vigorous to relatively
during subsequent events (for a more thorough review see, weak, and wax gourd is medium to weak. Most combinations of
Andrews and Marquez, 1993). Living cells from both root- watermelon and bottle gourd are acceptable, but a weak bottle
stock and scion then extend into the necrotic zone. A callus gourd rootstock was recommended by Hagihara (2004) when
bridge of interlacing parenchyma cells formed when cell di- using an extremely vigorous watermelon cultivar.
vision ruptures and invades the necrotic layer. During these A study by Edelstein et al. (2004) showed that number of
events the tensile strength of the graft increases due to phys- leaves, stem length, and fresh weight of melon plants increased
ical cohesion between rootstock and scion. New vascular cam- using 22 different Cucurbita spp. rootstocks. Davis and Perkins-
bium is differentiated from parenchyma cells, and secondary Veazie (2005) demonstrated that rootstocks affect the number of
xylem and phloem are produced by the reconstituted cam- nodes and lateral branches, and that the vigor of grafted water-
bium, providing a vascular connection between rootstock and melon plants was improved when grated onto a gourd rootstock.
scion. They further showed that grafting did not affect length, circum-
In some cucurbits, the stem (both the hypocotyl and the epi- ference, or diameter of fruit, but decreased weight of the fruit,
cotyl) has a central pit or cavity with 6 vascular bundles located suggesting the fruit were less dense.
in a definite position (Oda et al., 1993). Complete union of these
six vascular bundles is sometimes impossible depending on the
grafting methods used. Initial studies by Lu et al. (1996) indi- C. Yield and Size
cated that the addition of exogenous plant hormones helps the Grafting can increase yield since grafted plants are resistant
formation of graft unions by increasing the rate of formation to soilborne disease, have strong root systems, and increased
and number of vascular bridges. The differentiation, and pos- photosynthesis (Xu et al., 2005b; Qi et al., 2006; Wu et al.,
sibly the induction, of the vascular bridge in Cucumis sativus 2006). Since grafted plants are healthier and live longer, the har-
requires high levels of the auxin, indole-3-acetic acid (IAA), vesting period can be expanded dramatically. This advantage
which induces callus proliferation from the two partners at the is more obvious under less optimal growing conditions (Hang
graft union (Lu and Song 1999; Lu, 2000). Lu and Song (1999) et al., 2005). Cucumber plants grafted onto pumpkin rootstocks
mentioned that IAA and zeatin plus zeatin riboside are required had 27% more marketable fruit per plant than self-rooted cucum-
for vascular bundle regeneration in the graft union. There is a ber (Seong et al., 2003). Salam et al. (2002) demonstrated a 3.5
positive correlation between the vigor of grafted watermelon times higher yield due to larger fruit size, more fruit per plant,
and the similarity of protective isozymes (e.g., peroxidase and and improved plant survival rates. A squash interspecific hybrid
superoxide dismutase) between grafted and regular seedlings rootstock increased watermelon yield and increased fruit size
(Zhang et al., 2006b). (Miguel et al., 2004). Yetisir and Sari (2003) and Yetisir et al.
Studies by Ruiz and Romero (1999) demonstrated that grafted (2003) tested multiple rootstocks (Cucurbita moschata, Cucur-
melon plants had lower concentrations of nitrate, higher concen- bita maxima, squash interspecific hybrids, and bottle gourd) for
tration of nitrate reductase activity, and lower contents of total effects on yield. They found that watermelon scions grated onto
free amino acids and soluble proteins compared to the control. bottle gourd had 27–106% greater yield over control plants, but
Organic N and fruit yield also increased significantly in the ma- the Cucurbita sp. rootstock decreased yield by 127–240%. In
jority of grafted plants. These changes were attributed to dif- a review, Sakata et al. (2007) wrote that yield and fruit weight
ferences in N utilization and assimilation between grafted and of ‘Shin-tosa’ grafted watermelons were higher than those with
control plants. With these important physiological effects on the other rootstock. This was especially true in watermelon, and to a
scion, it is not surprising that grafting affects scion vigor, yield, lesser extent in melon (Miguel et al., 2004). Higher yields were
and fruit quality. not noted with grafted cucumber (Brajeul and Letard, 1998).
To complicate the matter further, yield was not affected by root- opment (about 30% fruit growth) in seeded heirloom varieties
stock in most scion / rootstock combinations with Cucumis melo and in Cucumis melo (Elmstrom and Davis, 1981; Chisholm
scions (Koutsika-Sotiriou and Traka-Mavrona, 2002). It was and Picha, 1986; Hubbard et al., 1989), but was found to be
suggested that this discrepancy in the literature demonstrated present at all stages of growth in grafted and non-grafted wa-
the importance of optimizing rootstock / scion combinations for termelons (Liu et al., 2006). Acid invertase activity was high
each cropping environment (Ruiz et al., 1997). during the first third of watermelon fruit development, and then
In Cucurbita spp., yield is dependent mainly on plant vigor. had little activity during the last 5 days of ripening. Sucrose
This may be why yield of ‘Shin-tosa’ grafted watermelon are synthase activity was moderately high in the first third of fruit
higher than other combinations (Sakata et al., 2007). Yield of development, then remained low. Sucrose phosphate synthase
plants with weak rootstocks are inferior to the yield of those increased during the second third of fruit development, and fell
with vigorous ones. Even if plants with weak rootstocks are in the last five days of ripening. The low sugar content of grafted
spaced widely and grown in fertile conditions, their yields are watermelon is correlated with low invertase, high sucrose syn-
still low, due to few fruit per plant. In some cultivars, larger fruit thase activities, and low sucrose phosphate synthase activity,
size caused marketability problems, mainly in ‘Galia’ (Cucumis and sugar transmembrane transportation capability (Qian et al.,
melo var. saccharinus Naud.) type melons (Miguel, 2004). 2004; Liu et al., 2006). The increase of sucrose in grafted plants
The yield of bottle gourd-grafted watermelon is relatively was accompanied by an increase of sucrose phosphate synthase
stable, as it is affected little by soil characteristics (Tamada, and sucrose synthase activities (Xu et al., 2006a). In addition,
1989). Wax gourd-grafted and watermelon-grafted watermelon it appears that sugar accumulation in mature fruit varies with
are affected by soil conditions, and often show decreased yield rootstocks (Gao and Liao 2006; Xu et al., 2006b).
in areas of low fertility or moisture (Sakata et al., 2007). Melon: Preharvest internal decay or internal breakdown and
abnormal fruit fermentation occurred in oriental melons and
melons when grafted onto vigorous squash interspecific hy-
D. Quality brid (Muramatsu, 1981; Chung, 1995). Other abnormalities ob-
Changes in grafted cucurbit fruit quality appear to be both served were reduced fruit soluble solids content and persistent
rootstock, and scion-dependent, causing contradictory reports green color in the suture stripe (Lee, 1989; Lee et al., 1998).
in the literature. Additionally, as watermelon and muskmelon Cucurbita spp. rootstock sometimes negatively affected melon
(Cucumis melo var. reticulatus) maturity is often delayed in fruit quality (Brix, taste, texture) (Traka-Mavrona et al., 2000).
grafted fruit, it is important to sample the fruit several days Although ‘Shin-tosa’ rootstock is resistant to fusarium wilt, it
past the expected fully ripe date to make sure values for sug- is almost never used for melon in Japan because it causes re-
ars do not simply represent delayed ripening (Miguel, 1997; Xu duced fruit quality, including low sugar content, and fibrous flesh
et al., 2005d). This difference in maturation probably causes (Muramatsu, 1981). An off-taste and texture in winter melon
additional contradictory reports in the literature. fruit grown in Greece in some of rootstock/scion combinations
As early as the late 1940s, Imazu (1949) noted that Cu- were reported by Koutsika-Sotiriou and Traka-Mavrona (2002).
curbita moschata rootstock caused inferior texture and flavor If Cucurbita spp. are used as a rootstock with ‘Earl’s Favorite’
in grafted ‘Honey Dew’ fruits, despite conferring resistance to melon, the fruit grow better than those on nongrafted plants, but
fusarium wilt. Grafting reportedly caused a small reduction, ap- the quality of the fruit netting is poorer and the sugar content is 2
proximately 1◦ Brix, in the sugar content of both watermelon and to 3◦ Brix less (Kamiya and Tamura, 1964). Trionfetti-Nisini et
melon (Taussig et al., 1996; Miguel, 1997; Lopez-Galarza et al., al. (2002) reported that of 13 commercial melon rootstocks and
2004; Xu et al., 2005d; Qi et al., 2006; Xu et al., 2006b). Nev- various Cucurbitaceae spp. rootstocks, many of them influenced
ertheless, when grown properly, fruits from grafted plants were quality (shape, size, Brix). Grafting was reported to increase vit-
commercially marketable, and reduction in fruit quality was not rescence (glassy look to flesh) in melon (Taussig et al., 1996).
found in the Spanish type melons or cucumber (Miguel, 1997; The photosynthesis rate of grafted melon plants decreases dra-
Hoyos, 2001). matically during late stages of fruit development (Xu et al.,
Fortunately, under the best circumstances, and when using 2005a). This decrease of photosynthesis may be related to low
specific rootstock/scion combinations, grafting can enhance fruit fruit quality.
quality (total soluble solids, and carotenoid and ascorbic acid Watermelon: The most common complaints about grafted
contents) (Davis and Perkins-Veazie, 2005; Xu et al., 2005b; watermelon quality are low Brix, insipid taste, increased number
Alan et al., 2007; Alexopoulos et al., 2007). Breeding for im- of yellowish bands in the red flesh, and internal flesh breakdown
proved rootstock/scion combinations and screening for combi- (Ryu et al., 1973; Lee and Oda, 2003; Davis et al., 2008). Tex-
nations that result in high quality fruit, is beginning to decrease tural quality and firmness were lower in watermelon grafted
the number of negative quality issues for grafted cucurbit crops onto ‘Shin-tosa’ than those grafted onto bottle gourd or con-
(Davis et al., 2008). trol (Yamasaki et al., 1994). The deterioration of watermelon
The primary sugars in watermelon are sucrose, fructose, and fruit quality on Cucurbita spp. rootstocks was attributed to vig-
glucose. Sucrose appears only after initiation of color devel- orous nutritional uptake (Kato and Ogiwara, 1978; Shinbori
et al., 1981; Masuda et al., 1986). In heated greenhouse ex- fruits from grafted plants tend to be shorter and show decreased
periments, grafted watermelon on five rootstocks (bottle gourd firmness and shelf life (Lee, 1989; Lee et al., 1999).
and Cucurbita pepo) were more vigorous and out yielded the
non-grafted controls by 25 to 95% (Ioannou et al., 2002). It was
also found that fruit quality [external and internal appearance, IV. WHAT CUCURBITS ARE GRAFTED
intensity of red, presence of fibrous tissue, cracks, cavities, ab- A. Watermelon
normalities in the rind and organoleptic index (reflecting taste), Watermelon is an important crop in Asian countries. The
texture, and Brix] was superior in nongrafted plants. A complex total area harvested in China was 2.2 million hectares in 2005
study on the effect of grafting and cytokinin-induced fruit set- (FAOSTAT, 2005). This counts for more than half the world
ting on color and sugar development in triploid watermelon was watermelon production. In 2005, Korea ranked ninth in total
reported by Liao and Lin (1996). They demonstrated that de- world production, about 850,000 metric tons. In Spain all the
spite the loss of other quality traits, fruit from grafted plants set triploid cultivars and diploid pollinizer are grafted (Camacho
with CPPU had a total Brix content similar to fruit from control and Fernandez, 2000; Hoyos, 2004; Miguel, 2004). Watermelon
plants. is the most commonly grafted cucurbit in Italy (Amadio, 2007).
Quality (Brix, lycopene, flesh firmness, rind thickness, and Watermelon was first grafted onto Cucurbita moschata and
fruit shape) of watermelon was greatly affected by grafting shortly afterwards onto bottle gourd to control soilborne dis-
(Yetisir and Sari, 2003; Yetisir et al., 2003; Davis and Perkins- eases caused by successive cropping (Ashita, 1927; Tateishi,
Veazie, 2005). Flesh firmness was increased slightly, lycopene 1927; Sakata et al., 2007). Remarkable yield increases of more
was typically higher, and Brix typically lower in watermelon than 200% derived from growing grafted plants under protected
fruit from grafted plants using gourd rootstocks (Davis and structures attracted the interest of many growers (Lee and Oda,
Perkins-Veazie, 2005). The total sugar content of watermelons 2003). As a result, the area for protected cultivation increased in
grafted onto bottle gourd rootstock was reported to be lower Japan, Korea, and China, and the demand for grafted seedlings
than in self-rooted watermelons (Yao et al., 2003; Qian et al., markedly increased despite their high cost. In fact, grafting tech-
2004; Liu et al., 2006). They reported that Cucurbita sp. root- niques are often developed with watermelon in mind.
stock decreased the quality of watermelon fruit, but that fruit Squash interspecific hybrids, bottle gourd, and Cucurbita spp.
from scion grafted onto bottle gourd differed only slightly from are often used as alternate rootstocks for watermelon to pre-
control fruit. Miguel et al. (2004) found no difference in solu- vent disease infection. However, rootstocks of Cucurbita spp.
ble solids concentrations of watermelon fruit from scions grafted often lower fruit quality (Kato and Ogiwara, 1978; Shinbori
onto a squash interspecific hybrid versus control, but Salam et al. et al., 1981; Masuda et al., 1986). Wild type watermelon is
(2002) showed a marked increase in watermelon TSS content used as a rootstock to prevent the occurrence of fusarium wilt
when grafted onto bottle gourd. Perkins-Veazie et al. (2008) in grafted plants. It has a good graft compatibility with water-
demonstrated that grafting watermelon could increase lycopene melon, leading to increased fruit quality (Huh et al., 2002), but it
and total carotenoids by 20%, and increase amino acids, es- is susceptible to low temperature and high moisture injury. Burr
pecially citrulline [a nonessential amino acid with vasodilation cucumber (Sicyos angulatus L.) is fairly resistant to wilt disease
properties (Lee et al., 1996)], by up to 35%. and adaptable during low temperature, but it has poor and slow
Cucumis sativus and Cucurbita pepo: For crops harvested seed germination, making grafting difficult and inefficient (Lee
at an immature stage such as cucumber, grafting has few re- et al., 1999). Wax gourd has good graft compatibility with wa-
ported negative effects on fruit quality, although increased firm- termelon but is susceptible to low temperature stress that can
ness and shortening of fruit have been noted (Muramatsu, 1981). delay development and blooming period of the grafted plants
As mentioned above, ‘Bloomless’ cucumber fruits have gained (Kurada, 1974; Ko, 1999; Huh, 2000; Sakata, 2007).
a prominent market-share. However Sakata et al. (2008) report
inconsistency in the literature on ‘Bloomless’ cucumber tex-
ture. Some reports show they have less desirable texture due B. Cucumber
to tougher skin and softer flesh (Inayama, 1993a,b; Yamamoto Cucumber is a popular vegetable in China with over 1.7 mil-
et al., 1996). Questions remain as to whether traditional cucum- lion hectares of cucumber and gherkins harvested in 2005 (FAO-
bers taste better than ‘Bloomless’ fruits (Horie et al., 2004). STAT, 2005). About 50% of China’s cucumbers are grown in
The main reason for selecting rootstocks for cucumber culti- plastic tunnels during the spring and fall season, and in solar
vars is to increase yield, produce ‘Bloomless’ fruit, and achieve greenhouses during winter months in Northern China. Almost
stable production. However, it has been suggested that differ- all cucumbers grown in solar greenhouse are grafted.
ent rootstocks affect grafted cucumber quality characteristics Growing cucumbers under protected environments started in
such as fruit shape, skin and flesh color and texture, firmness, the 1960s with the introduction of commercial plastic films,
rind thickness, and ascorbic acid and soluble solids content enabling year-round production of several horticultural crops.
(Inayama, 1989; Choi et al., 1992; Kang et al., 1992; Robinson Compared to watermelons and melons, cucumbers do not man-
and Decker-Walters, 1997; Zhu et al., 2006b). Summer squash ifest an extensive spread of soilborne diseases, especially when
grown outdoors in the summer. Cucumber cultivars developed One of the most serious problems in greenhouse production
for winter greenhouses are more susceptible to various dis- of oriental melon is nematode control (Ko, 1999). Rootstocks
eases and unfavorable environmental conditions than those in- with good to excellent nematode resistance are required for suc-
tended for open field cultivation. Other advantages of grafting cessive melon production in many areas of Korea. Limited suc-
are low temperature tolerance, yield increase, ease of manage- cess was obtained with ‘Andong,’ a burr cucumber rootstock.
ment, higher fruit quality, ‘Bloomless’ fruit, and extension of Finding alternative rootstocks for nematode control is an im-
harvest period in hydroponic systems (Asao et al., 1999). portant area of research and burr and African horned cucum-
Cucurbita spp. and figleaf gourd are commonly used as root- bers (Cucumis metuliferus E. Meyer ex Naudin) are considered
stocks for cucumber. Figleaf gourd is by far the most popular of promising (Igarashi et al., 1987). Grafted melons in Spain are
the two species. mostly ‘Galia,’ cantaloupe, and Armenian cucumber (Cucumis
Figleaf gourd is a common rootstock for cucumber since melo var. flexuosus Naud) types (Miguel, 2004). Cantaloupe
the two species are highly compatible. This rootstock pos- types are grafted in France and Italy (Taussig et al., 1996; Buzi
sesses good to high fusarium wilt (Ko, 1999), phytophthra, root et al., 2002).
knot nematode resistance and low soil temperature tolerance
(Marukawa and Takatsu, 1969a; Ahn et al., 1999; Lee and Oda, D. Bitter Gourd
2003). It also has good water and nutrients absorption (Shimada Bitter gourd is native to India and is grown throughout Asia.
Both immature fruit and vine tips are nutritious and edible. The
and Moritani, 1977; Masuda and Gomi, 1984) even at low tem-
peratures (Tachibana 1987; 1989). The disadvantage of figleaf main problem with bitter gourd production is fusarium wilt and
as a rootstock is its rapid seedling growth, making the optimal low temperatures during winter production (Lin et al., 1998).
grafting window short. In addition, the germination rate is low Luffa (Luffa spp.), figleaf gourd, and pumpkin are common root-
and not uniform (Hang et al., 2005). stocks for bitter gourd (Lin, 2004). Luffa has been widely used
Squash interspecific hybrid rootstocks are more tolerant to during summer production since it is resistant to fusarium wilt
high temperature conditions and are commonly used for cu- and has good tolerance to heat and flooding. Figleaf gourd and
cumber during summer plantings. Cucumber grafted onto cer- pumpkin are used by farmers during low temperature winter pro-
tain cultigens of Cucurbita moschata produced fruits with shiny duction. Since figleaf gourd does not have good heat tolerance,
skin compared to fruit from nongrafted plants or from cucum- bitter gourd-figleaf grafted plants often show premature decay
ber grafted onto other Cucurbita spp. (Fujieda, 1994). The shiny during hot weather. To overcome this problem, figleaf gourd and
‘Bloomless’ fruits are successful in the Japanese market, but not luffa, are often used together. Figleaf gourd roots will support
in Korea. In France, squash interspecific hybrids or Cucurbita fi- the scion early and as temperatures rise, luffa roots support the
cifolia are used as rootstocks for cucumber (Brajeul and Letard, plant. Cleft grafting is a good method for grafting bitter gourd
1998). In Spain the best rootstocks for cucumber have been the (Lin, 2004).
squash interspecific hybrids (Hoyos, 2001). E. Summer Squash
Burr cucumber is fairly resistant to wilt disease and adaptable Summer squash is grown widely under protected culture
during low temperature. However, it has poor and slow seed during winter in China. Grafted plants are more compact and
germination, making grafting difficult and inefficient (Takeuchi tolerant to low soil temperature, and fruit are uniform and
and Nagai, 1977; Hagitani and Toki, 1978; Toki, 1978). straight. This fruit shape is preferred by Chinese customers. The
most commonly used rootstocks are figleaf gourd and Cucurbita
C. Melons
spp. (Hang et al., 2005).
In Spain and Italy, all grafted melon plants are now on squash
interspecific hybrids, but B. hispida or Cucurbita maxima was
used in commercial production before the introduction of the F. Chinese Snake Gourd
hybrids (Bianco, 1990; Chaux and Foury, 1994; Messiaen et al., Chinese snake gourd (Trichosanthes kirilouii Maxim) is a
1994; Miguel and Maroto, 1996; Fritsch, 2002). In Italy, cultivars dioecious crop. Female and male plants are indistinguishable
of Cucurbita maxima tolerant to Monosporascus vine decline before transplanting. Farmers sometimes use male plants for
(Gennari, 1999) are used except in places where this disease is rootstocks and female shoots for scions to increase fruit yield
not prevalent. Locations where Monosporascus vine decline is (Li et al., 2006). Grafting onto wax gourd improves fusarium
not an issue, Cucumis melo lines resistant to Fusarium oxyspo- wilt resistance (Lü et al., 2004).
rum f. sp. melonis are used (Buzi et al., 2004).
Grafting is a must in oriental melon cultivation since their G. Commonly Used Rootstocks
roots are sensitive to low soil temperatures. However, grafting Tables 4 and 5 demonstrate the various uses of different root-
onto vigorous rootstocks induces undesirable physiological dis- stocks. Marukawa and Yamamuro (1967) examined 84 cultivars
orders such as vigorous vegetative growth, delayed fruit maturity and accessions of Cucurbita spp. for use as potential water-
and harvesting, uneven maturity, and internal fruit breakdown melon rootstocks. They concluded the following: 1) about 20%
(Chang, 1995). of the tested lines of Cucurbita maxima, Cucurbita moschata,
TABLE 4
Response of cucurbits to biological and environmental stresses
Nematode Graft compatibility
Fusarium
Rootstock M. M. Low temp High salt Oriental
and scion Ia II III IV incognita halpa tolerance tolerance Watermelon Cucumber melon
Rootstockb
Shintozwa HRc HR HR HR S S HR HR HCd HC HC
Hongtozwa HR HR HR SR S S MR MR SC HC HC
Figleaf gourd MR SR MR SR S S HR HR IC HC IC
Bottle gourd MR HR HR SR S S SR MR HC HC IC
Wax gourd HR MR HR HR S SR SR SR HC HC —
Bur cucumber HR HR HR HR S HR SR SR HC MC HC
AH cucumbere HR HR HR HR S MR SR ? HR HC HC
Scion
Watermelon S SR HR HR HR SR S SR — — —
Cucumber HR SR HR HR S S HR SR — — —
Oriental melon HR HR S HR S S S S — — —
a
I, Fusarium oxysporum f. sp. niveum II; F. oxysporum f. sp. cucumerinum; III, F. oxysporum f. sp. melonis; and IV, F. oxysporum f. sp.
lagenariae.
b
Shintozwa (Cucurbita maxima x Cucurbita moschata), Hongtozwa (Cucurbita moschata), figleaf gourd (Cucurbita ficifolia), bottle gourd
(Lagenaria siceraria), wax gourd (Benincasa hispida), bur cucumber (Sicyos angulatus), and AH cucumber (Cucumis metuliferus), respectively.
c
HR, highly resistant; MR, moderately resistant; SR, slightly resistant; and S, susceptible.
d
HC, highly compatible; MC, moderately compatible; SC, slightly compatible; and IC, incompatible.
e
AH: African horned cucumber.
Source: Ko (1999).
and Cucurbita pepo were compatible with watermelon; 2) figleaf V. GRAFTING METHODS
gourd was incompatible with watermelon; and 3) the accessions The survival rate of grafted plants depends on compatibility
‘Kinshi Uri’ (Cucurbita pepo), ‘Bizen Chirimen,’ and ‘No. 8’ between scion and rootstock, quality and age of seedlings,
(Cucurbita moschata) were suitable rootstocks for watermelon. quality of the joined section, and post-grafting management.
The squash interspecific hybrids are the most commonly used Different grafting techniques are adapted for different scions
cucurbit rootstocks, but some cultivars of Cucurbita moschata, and rootstocks depending on grafting objectives, farmers’ expe-
Cucurbita pepo, and Citrullus spp. are increasing in importance rience, and post-grafting management conditions. When scion
(Miguel and Maroto; 1996; Paplomatas et al., 2002; Miguel, and rootstock have hollow hypocotyls, the hole insertion and one
2004). Bottle gourd and wax gourd are suitable rootstocks for cotyledon grafting methods are preferred (Hang et al., 2005).
watermelon because they are resistant to fusarium wilt, have high In contrast, tongue approach and cleft techniques, which have
affinity for watermelon, and they provide stable plant growth high survival rates, are often chosen by inexperienced farmers
after grafting (Sato and Takamatsu, 1930; Matsumoto, 1931; who have plenty of space and adequate labor. In contrast, the
Tateishi, 1931; Kijima, 1933; Murata and Ohara, 1936; Kijima, one cotyledon and the hole insertion grafts require experienced
1938). Bottle gourd is not used in Spain because of its suscep- labor, specialized tools, and a healing chamber to obtain high
tibility to monosporascus vine decline (Garcı́a-Jiménez et al., survival rates. Cleft, one cotyledon, and hole insertion methods,
1994), but in Italy it is used as well as Cucurbita moschata which have high grafting positions, increase separation of
and squash interspecific hybrids (Morra, 1997; Buzi et al., scion from the ground. This decreases the opportunity of scion
2004). adventitious roots contracting soilborne diseases (Hang et al.,
Burr cucumber, which shows fusarium wilt and nematode 2005).
resistance, low temperature tolerance, and tolerance of excess In manual grafting, the grafting and post-grafting operations
water, was tested extensively for use as a rootstock for cucum- require three to four people, each assigned to a specific step in
ber (Hagitani and Toki, 1978; Toki, 1978). However, it was not the process (Lee and Oda, 2003). Automation requires one to
adopted because of its susceptibility to damping-off and gummy three people to oversee the entire process. Below is a review
stem blight (Takeuchi and Nagai, 1977), uneven germination, of the published grafting methods, both manual and automated,
and its thin hypocotyl makes grafting difficult (Tataki, 2004). and their pre- and post-grafting requirements.
TABLE 5
Rootstocks for cucurbitaceous crops and some related characteristics
Scion/Rootstock Cultivar Major characteristics Possible disadvantage
Watermelon
Bottle gourd (Lagenaria FR Dantos, Partner, Vigorous root system, resistant to New fusarium race,
siceraria) Renshi, FR Combi, fusarium and low temperature susceptible
TanTan anthracnose
Squash (Cucurbita Chinkyo, No. 8, Keumkang Vigorous root system, resistant to Poor fruit shape and
moschata) fusarium and low temperature quality
Interspecific hybrid squash Shintozwa, Shintozwa #1, Vigorous root system, resistant to Reduced fertigation
(Cucurbita maxima × C. Shintozwa #2, Chulgap fusarium and low temperature required; reduced
moschata) excellent vigor and high quality
temperature tolerance
Pumpkins (Cucurbita pepo) Keumsakwa, Unyong, Vigorous root system, resistant to Poor fruit shape and
Super Unyong fusarium and low temperature quality
Wintermelon (Benincasa Lion, Best, Donga Good disease resistance Incompatibility

hispida)
Watermelon (Citrullus Kanggang, Res. #1, Fusarium tolerance, but not Not enough vigor and
lanatus) Tuffness, Kyohgoh. resistance disease resistance
African horned cucumber NHRI-1 Excellent fusarium resistance and Medium to poor graft
(Cucumis metuliferus) good nematode tolerance compatibility
Cucumbers
Figleaf gourd (Cucurbita Heukjong, Black Seeded, Good low temperature tolerance Fruit quality may be
ficifolia) Figleaf gourd and disease resistance reduced
Squash (Cucurbita Butternut, Unyong #1, Good fusarium tolerance and Affected
moschata) Super Unyong ‘bloomless’ fruit skin byPhytophthora
Interspecific hybrid squash Shintozwa, Keumtozwa, Good fusarium and low Slight quality reduction
(Cucurbita maxima x C. Ferro RZ, 64-05 RZ, temperature tolerance expected
moschata) Gangryuk Shinwha,
Chulgap
Bur cucumber (Sicyos Andong Good fusarium tolerance, low and Reduced yield
angulatus) high soil moisture tolerance and
nematode tolerance
African horned cucumber NHRI-1 Excellent fusarium resistance and Weak temperature
(Cucumis metuliferus) good nematode tolerance tolerance
Melons-Oriental Melons
Squash (Cucurbita Baekkukzwa, No. 8, Good fusarium and low Phytophthora infection
moschata) Keumkang, Hongtozwa temperature tolerance
Interspecific hybrid squash Shintozwa, Shintozwa #1, Good fusarium resistance, low and Phytophthora infection,
(Cucubita maxima x C. Shintozwa #2 high soil temperature tolerance, poor fruit quality
moschata) and high soil moisture tolerance
Pumpkin (Cucurbita pepo) Keumsakwa, Unyong, Good fusarium resistance, low and Phytophthora infection
Super Unyong high soil temperature tolerance,
and high soil moisture tolerance
Melon (Cucumis melo) Rootstock #1, Kangyoung, Fusarium tolerance and good fruit Phytophthora problem
Keonkak, Keumgang quality
African horned cucumber NHRI-1 Good fusarium tolerance, low and Weak temperature
(Cucumis metuliferus) high soil moisture tolerance and tolerance
nematode tolerance
Source: Asao et al. (1999); Chung (1995); Igarashi, Kanno, and Kawaide (1987); Gomi and Masuda (1981); Kang, Choi, and Lee, 1992; Kim and
Lee (1989); Ko (1999); Matsuo, Ichiuchi, and Kohyama (1985); Lee (1989; 1994; 2000); Lee et al. (1998); Oda (1999); Oh (2000); Tachibana
(1981).
TABLE 6
Efficiency and savings of labor by using three different
grafting methods
Labor Labor saved
Treatment (min/100 plants) (%)
PG 46.0 35.8
HIG 60.3 15.6
TAG 71.4 0
PG (Pin grafting), HIG (Hole insertion grafting), TAG (Tongue ap-
proach grafting).
Methods continue to evolve to overcome existing and devel-

oping problems, to increase survival rate, and decrease labor. The
following methods are summarized from Hassell and Memmott
(2008), Kapiel et al. (2004), Lee (1994), Lee et al. (1998), Lee
and Oda (2003), and Oda (1999). Table 6 shows the efficiency
of three of the main grafting techniques.
A. Tongue Approach/Approach Graft

The advantage of the tongue approach, which is sometimes
called approach graft (TAG), is that it is easy and requires a low FIG. 1. The tongue approach grafting method. Step 1, preparing the rootstock;
step 2, preparing the scion; step 3, joining the scion to the rootstock; step 4,
relative humidity microclimate after grafting. Because of this,
securing the joining with a metal strip; and step 5, removing the scion roots.
it is often adapted by farmers. Although this method requires Reprinted with permission from HortScience (Hassell and Memmott, 2008).
more space and labor compared to other methods, high seedling
survival rate can be attained even by beginners. However, the
ing. If the top of the rootstock was left on, it should be re-
grafting position is close to the ground making it easy for ad-
moved five days after grafting. The scion hypocotyls are cut off
ventitious roots from the scion to reach the soil. Moreover, this
seven to ten days after grafting at just below the graft union.
method is not suitable for rootstocks with hollow hypocotyls,
Grafted plants are maintained in the greenhouse until ready for
since scions may form adventitious roots inside the hypocotyls.
transplanting. This is at least two days after removing the scion
Therefore the TAG method is good for rootstocks with high solid
roots. A humidity chamber is not required (Lee, 1994; Lee et
hypocotyls.
al., 1998; Oda, 1999; Lee and Oda, 2003; Hassell and Memmott,
This method originated in the Netherlands (Ishibashi, 1965),
2008).
and is now widespread in Japan, Spain, France, and Italy
(Bianco, 1990; Morra, 1997; Brajeul and Letard, 1998; Miguel
and Maroto, 2000; Buzi et al., 2002; Lee and Oda, 2003). It is B. Hole Insertion/Terminal/Top Insertion Graft
easy to use, has a high success rate, and the grafted seedlings The hole insertion graft (HIG), also known as terminal graft
have a uniform growth rate. and top insertion graft, is the most popular cucurbit grafting
The scions and rootstocks should be approximately the same method in China. This method is easy, has high survival rates,
diameter in the TAG method. This is usually the case after and the grafted plants have fewer incidents of soilborne disease
the rootstock has fully developed cotyledons and the scion has because of the high graft union (Xiao and Yan, 2004; Hang
cotyledons and the first true leaf. The rootstock is cut through et al. 2005). One person can produce 1,500 or more grafts/day
the hypocotyl at a 35◦ to 45◦ angle (Figure 1). The growing and post-grafting acclimatization is simple. This method was
point may remain by only cutting downward halfway through described in 1970 by Fujii and has recently been adopted by
the hypocotyl, or it may be removed with a grafting knife or nurseries (Sakata et al., 2007). A modified HIG method, where
razorblade. The scion is also cut at an angle (upward if the the root system is removed from the rootstock, was originally
growing tip of the rootstock remains attached) and joined to developed for watermelon grafting (Utsugi, 1973) and will be
the rootstock with the cut surfaces aligned. Lead strips, alu- discussed below under ‘Root Pruning.’
minum foil, or grafting clips are used to hold the grafts together. Normally rootstock seed are sown 2–4 days before scion
When metal strips are used they can remain on the plant once seeds. Rootstocks are ready for grafting when both cotyledons
the plant has healed. However, the grafting clips need to be and first true leaf start to develop (about 7 to 10 days after sow-
removed once the union is healed. There is an additional cost ing). The rootstock’s growing point is removed (Figure 2), a cut
for clips, and for labor to remove them 15–20 days after graft- is made and then drilled at a 35◦ to 45◦ angle using a bamboo
FIG. 3. The one cotyledon grafting method. Step 1, preparing the scion; step
2, preparing the rootstock; step 3, joining the plants; step 4, securing the joined
region with a grafting clip. Reprinted with permission from HortScience (Hassell
and Memmott, 2008).
Sakata et al., 2007). This method is suitable when rootstock and

scion are of a similar size. The rootstock should be grafted when
cotyledons and the first true leaf start to develop (about 7 to 10

days after sowing). One cotyledon and the growing tip are re-
moved (Figure 3). The seedling is cut at a slant from the base of
one cotyledon to 0.8–1.0 cm below the other cotyledon, remov-
ing one cotyledon and the growing tip. The length of cut on the
scion hypocotyl should match that of the rootstock and should
be at a 35◦ to 45◦ angle. The scion is attached to the rootstock
FIG. 2. The hole insertion grafting method. Step 1, preparation of the scion;
step 2, removing the growing tip of the rootstock; step 3, drilling out the growing and fixed tightly by a grafting tube or clip (Oda, 1999; Lee and
tip and creating a hole for the scion; step 4, joining the scion and rootstock; step Oda, 2003). Grafted plants should be maintained in the dark at
5, the joined plants. Reprinted with permission from HortScience (Hassell and 25◦ C and 100% humidity for three days or until the junction
Memmott, 2008). has healed, before moving them into a greenhouse maintained
at 21◦ C to 30◦ C. Plants should not be more than 33 days old
or plastic gimlet. The scion hypocotyl is cut to form a 7–8 mil- before transplanting (Lee, 1994; Lee et al., 1998; Kapeil et al.,
limeter long wedge. The scion is then inserted into the prepared 2004). This approach requires good post-grafting management
rootstock hole. This method does not require additional labor since scions easily fall off at an early stage. OCG is called tube
for clipping, transplanting, cutting off remaining scion roots, or grating when the joined plants are held together with a length of
clip removal. It does require a higher skill level compared to tube instead of a grafting clip (Hassell and Memmott, 2008).
TAG, and requires a suitable environmentally controlled humid
or healing chamber. D. Cleft/Side Insertion Graft
To achieve a high rate of success, relative humidity should be The cleft graft (CG) also known as the side insertion graft
maintained at approximately 95%. After the healing process is was the initial grafting method used for cucumber and water-
complete, grafted plants are transferred and maintained at 21– melon (Ishibashi, 1959). The CG has largely been replaced by
36◦ C in the greenhouse until junction is healed. Plants should OCG, HIG, and TAG, because of the higher success rate and the
not be older than 33 days before transplanting (Lee, 1994; Lee uniform growth obtained from those methods (Amadio, 2004).
et al., 1998; Lee and Oda, 2003; Kapeil et al., 2004; Hassell and However, in Italy and France CG is still one of the most com-
Memmott, 2008). The main problem with HIG is that the size mon techniques (Bianco, 1990; Morra, 1997; Brajeul and Letard,
of the rootstock hole is limited by rootstock size. Therefore, the 1998; Buzi et al., 2002).
scion size has to be controlled, narrowing the grafting period. The CG method is suitable for rootstocks with wide
hypocotyls. This method is simple and easy to learn, and is suit-
C. One Cotyledon/Slant/Splice/Tube Graft able for preventing soilborne diseases since the grafting junction
The one cotyledon graft (OCG) also known as slant, splice, or is high on the hypocotyl. When cotyledons and the first true leaf
tube graft is commonly used, and has recently been adopted by start to develop (about 7 to 10 days after sowing) the rootstock is
commercial plug seedling nurseries (Fujii, 1970, Sakata et al., ready to graft. The growing tip of the rootstock can be removed
2007). The procedure is performed by hand, machine, or robot, during grafting or around five days after the grafted plants are
and is applicable to most vegetables. This approach is pre- removed from their high humidity chamber (Figure 4). A slit
ferred when rootstocks have thin stems, i.e. watermelon, cu- is cut on the rootstock hypocotyl with a razor blade and held
cumbers, and oriental melons (Yoshioka, 2001; Amadio, 2004; open with a toothpick. The width of the cut varies depending
FIG. 4. The cleft grafting method. Step 1, preparing the scion; step 2, preparing the rootstock; step 3, opening up the rootstock hypocotyl; step 4, inserting the
scion; step 5, securing the joined plants with a grafting clip. Reprinted with permission from HortScience (Hassell and Memmott, 2008).
on the diameter of the scion’s hypocotyl. The scion hypocotyl F. Double Graft
is cut from both sides at a 35◦ to 45◦ angle and then inserted Double grafting is used when a perfect rootstock for a specific
into the slit in the rootstock. The toothpick is removed. Then the scion is unavailable. For example, if the rootstock is too large
two cut surfaces are matched and held together with a grafting and an intermediate sized stem is needed to bridge the rootstock
clip or silicone sleeve. Grafted plants should be transferred to and scion. The scion is first grafted onto a middle rootstock (Fig.
a humidity chamber or healing room. Plants should be main- 5). The middle rootstock is then grafted onto another rootstock,
tained in the greenhouse until the junction is healed, and should which has good resistance to soilborne diseases. Unfortunately
not be more than 33 days old before transplanting (Lee, 1994; this method increases labor and cost while decreasing survival
Lee et al., 1998; Oda, 1999; Lee and Oda, 2003; Hassell and rate (Lee et al., 1998).
Memmott, 2008).
G. Root Pruning
E. Pin Graft Root pruning hole insertion grafting, and root pruning one
cotyledon grafting employ the same procedures as the HIG and
This method is similar to OCG, except that specially designed
OCG methods except the rootstock hypocotyl is cut to remove
pins are used instead of grafting clips to secure the graft union.
the roots. This induces adventitious root growth, increases the
The cotyledons of the rootstock and scion are cut horizontally,
production of primary roots, and enhances the plant’s tolerance
and a ceramic pin is inserted into the cut surface (Figure 5). This
to cold and heat thus ensuring vigorous growth (Lee and Oda,
helps align and secure the joined sections. This method’s limiting
2003). At present, more than 40% of watermelon grafting in
factor is that the scion and rootstock should be approximately
Japan is performed with this method. However, this added pro-
the same diameter so the cambial regions are in close contact.
cedure necessitates an exclusive grafting facility and is more
This method is easy, reducing labor cost, but ceramic pins add
dependent on soil conditions (Yoshida and Harada, 1972; Lee
expense, and a special environmentally-controlled chamber is
et al., 2001b; Xu et al. 2002, Zhu et al. 2006a).
needed to acclimatize the grafted plants (Lee et al., 1998).
H. Automated Graft
The first semiautomatic cucumber grafting system was com-
mercialized in 1993 and others have been developed since. A
simple grafting machine can produce 350–600 grafts per hour
with 2 operators, whereas manual grafting techniques produce
about 1,000 grafts per person per day (Masanao, 1996; Suzuki
et al., 1998; Lee and Oda, 2003; Gu, 2006). That is an increase
of 400 to 3,000 grafted plants per person per day.
In Spain, automated methods represent less than 5% of the
total cucurbit grafts, but is much higher in Japan. Grafting robots
are being developed in Japan and Korea that are more forgiving
or seedling uniformity and require less labor to operate than
older grafting machines. They are expensive and require highly
FIG. 5. Demonstration of Pin Grafting (1) and Double Grafting (2). uniform germination. The robots typically use the root pruning
OCG method (Masanao and Hisaya, 1996; Suzuki et al., 1998; ide) to minimize possible microorganism damage (Hassell and
Tiezhon and Liming, 2002; Lee and Oda, 2003). Kubota et al. Memmott, 2008).
(2008) report a fully automated grafting robot for cucurbits with
scion and rootstock feeders that pick, orient, and feed the scion
K. Compatibility
and the rootstock shoots to the grafting processor, performing
As grafting is a composition of two different plants, some
750 grafts per hour with a 90% success rate. The new robot will
beneficial or negative effects (besides the effect of soilborne
be commercially available in a few years.
pathogens), may arise after grafting. Graft incompatibility, as re-
viewed by Andrews and Marquez (1993), is differentiated from
I. Acclimatization graft failure that often results from environmental factors or lack
Proper environmental conditions are important to facilitate of skill of the grafter. When grafting conditions have been suc-
rootstock and scion union for some grafting methods. Some cessfully ensured, graft incompatibility could be attributed to
nurseries have chambers that maintain temperature above 20◦ C, other factors like failure of rootstock and scion to effect a strong
or for the tongue approach method, at around 25◦ C, and low union, failure of the grafted plant to grow, or premature death
light intensity for the first five to seven days. Relative humidity of either rootstock or scion after grafting. Also, physiological
should be maintained between 85% to 100% (Miguel, 1997). incompatibility may occur due to lack of cellular recognition,
Most nurseries acclimatize grafted plants in small plastic tun- wounding responses, presence of growth regulators, or incom-
nels inside greenhouses where it is possible to maintain a high patibility toxins.

relative humidity. Three to eight days after grafting, plants are Incompatibility induces undergrowth or overgrowth of scion
acclimated to the natural conditions of the greenhouse by slowly (Figure 6), which can lead to decreased water and nutrient flow
dropping humidity (Miguel, 1997; Lee and Oda, 2003; Hassell through the graft union and cause wilting of the plant. Generally,
and Memmott, 2008). grafting compatibility is related to taxonomic affinity but there
are significant exceptions. For example, luffa and melon have
J. Production of Rootstock and Scion higher compatibility with netted melon (Cucumis melo var. retic-
Cucurbits are usually grafted after the first true leaf appears, ulatus) compared to Chinese pumpkin (Cucurbita moschata)
but before it is fully developed in both the rootstock and scion and wax gourd (Wei et al., 2006). Grafting incompatibility usu-
seedlings. This means that planting dates must be optimized ally occurs at early stages as vascular connections are forming,
for both the scion and the rootstock. Uniformity in both ger- but incompatibility can appear as late as the fruiting stage, when
mination and growth of the rootstock and scion are critical for a high demand for nutrients and water is placed on the plant.
grafting survival. Optimum seedling age varies for species and Cucumber and trellised muskmelon plants grafted onto Cu-
different grafting methods. Difficult seeds to germinate, such as curbita rootstocks may collapse even in disease-free soils. This
triploid watermelon seeds, should be watered lightly and main- wilt was observed slightly more often in cucumber grafted onto
tained around 30◦ C for consistent germination rates (Hassell figleaf gourd (Yamamoto, 1979). This phenomenon appears dur-
and Schulthesis, 2002). If seedlings are too young, they are too ing the harvest period and was a serious issue in the 1970s in
delicate to handle during the grafting process, and if too old, Japan and Israel (Cohen et al., 2007; Sakata et al., 2008). In
may cause unwanted meristematic regrowth of rootstock tissue the major watermelon production areas, a similar sudden wilt of
(Hassell and Memmott, 2008). Some grafting facilities rinse leaf and stem occurred as early as the 1950s throughout Japan.
the seedlings with clean water, treated with fungicides or disin- Wilting occurred from the fruiting stages until harvest (Tamada,
fectant (e.g., Physan 20 or peroxyacetic acid/ hydrogen perox- 1989). It seems that this is a physiological disorder caused by
FIG. 6. Grafting compatibility between scion and rootstock.

an imbalance of growth between the scion and the rootstock VI. CONCLUSION
(Hamaya and Ogawa, 1973). Plants collapse because tylosis de- For decades, grafting has been successfully practiced in many
velops leading to a disrupted water supply and a rapid drop in Asian countries, and is becoming increasingly popular in Eu-
carbohydrates in shoots and roots as a result of a lack of solar ra- rope. Grafting is routinely employed for cucumbers, melons,
diation or severe pruning (Takahashi and Shiraki, 1976; Kondo, oriental melons, squash, and watermelon. Identification of com-
1978). Fruit load subjects plants to a progressive increase of patible multi-disease-resistant rootstocks with tolerance to abi-
water stress (Pivonia et al., 2002). Trellised grafted plants with otic stresses is a basic requirement for continued success. Ad-
high fruit load cannot always keep up with the water demand ditionally, the availability of efficient grafting machines, and
during hot periods. These plants are exposed to solar irradia- grafting robots will encourage cultivation of grafted cucurbits
tion from all directions, whereas prostrate plants are irradiated worldwide (Lee and Oda, 2003). Automatic grafting machines
only in their upper canopy (Cohen et al., 2007). To avoid this can increase grafting speed and increase the survival rate of
physiological sudden wilt, the promotion of root development grafted plants (Gu, 2006). They may also increase grafted cu-
and improved watering management is important. curbit crop production, and in turn, increase yield, and decrease
Watermelons do not have compatibility problems with the chemical usage.
usual rootstocks, mainly squash interspecific hybrids and Citrul- The low rate of grafting success results in high costs, reducing
lus spp. (Miguel, 2004). However, problems have been reported the appeal of grafting in developed countries. Increased graft-
with some rootstocks of Cucurbita moschata (Miguel, 1997) ing success requires close contact between scion and rootstock
and burr cucumber (Miguel et al., 2005). Yetisir and Sari (2003) vascular bundles, rootstock and scion compatibility, and proper
tested 10 rootstocks (Cucurbita moschata, Cucurbita maxima, environmental conditions to facilitate rootstock and scion union.
bottle gourd, and a squash interspecific hybrid). They found that Grafted plants with a good rootstock / scion combination have
the watermelon scion had a higher survival rate (95%) on bottle disease resistance, better abiotic stress tolerance, and higher
gourd type rootstock than on the Cucurbita spp. rootstock (65%). yield. Unfortunately, the range of commercial rootstocks is lim-
In melon, the cultivars of Spanish types (Cucumis melo ited, and the effect of unexplored rootstocks on fruit quality is
var.saccharinus) have serious compatibility problems with the still not clear.
squash interspecific hybrids (Miguel, 2004). However, the Several problems commonly associated with grafting have
‘Galia’ and Cantaloupe types, and some Cucumis melo var. flex- been reported such as: a) additional cost for rootstock seeds;
uosus (Miguel, 2004), have no compatibility problems with the b) labor for grafting and seedling maintenance; c) inadequate
squash interspecific hybrids. Morra (1997) demonstrated the low experience and technique for grafting and cultivation of grafted
affinity of melon with L. siceraria, Cucurbita ficifolia, and Cu- plants; and d) physiological disorders associated with grafting
curbita moschata but had good affinity with squash interspecific (Lee, 1994). Despite these problems, enormous benefits are real-
hybrids and Cucumis melo. ized from grafting, namely: a) avoidance of soilborne diseases;
Cucumber adapts well to grafting and has few compatibility b) added income from increased yield and premium price from
problems with the usual rootstocks (Hoyos, 2001). Oda et al. off-season production; c) lower cost of fertilizers and irrigation
(1993) performed experiments with horizontal grafting of cu- water due to the rootstocks extensive root systems; d) extended
cumber seedlings onto Cucurbita spp. rootstocks. Along with harvest periods; e) avoidance of soil salinity; f) reduced cost for
Yetisir and Sari (2004), they demonstrated that the survival rate soil fumigation; and g) a means to achieve organic vegetable
was inversely correlated with difference in the scion and root- production without fumigation (Lee and Oda, 2003). Further re-
stock hypocotyl diameters. The number of vascular bundles did search needs to focus on rootstock development, more efficient
not significantly affect survival rate, but did positively affect grafting robots, and development of acclimatization facilities.
the growth rate of grafted watermelon plants—in general, bot- This research should considerably reduce the cost of grafted
tle gourd type rootstocks had a higher survival rate than other seedling production in the future.
rootstocks. However, results from Edelstein et al. (2004) do not
concur. Their studies showed that stem diameter and number of
rootstock vascular bundles did not correlate with melon scion ACKNOWLEDGEMENTS
fresh plant weight with 22 Cucurbita spp. rootstocks. We would like to thank Amy Helms, Antoine Bastien van
Most cucurbit scions can be grafted onto several rootstock der Meer, Jim McCreight, Yun-Chan Huh, and Jung-Myung
species and incompatibility can be circumvented by using appro- Lee for valuable additional information and support. We also
priate grafting methods and growing environments (Ko, 1999). thank Richard Hassell and Amnon Levi for critical reviews of
Rojas and Rivero (2001) showed that grafting method and vari- the manuscript.
ety used significantly affected graft survivability, but scion age
(one to four weeks) did not. Moreover, Tamada (1969) pointed REFERENCES
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Critical Reviews in Plant Sciences

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Angela R. Davis,1 Penelope Perkins-Veazie,1 Yoshiteru Sakata,2

I. CUCURBIT GRAFTING ......................................................................................................................................51

II. WHY GRAFT CUCURBITS? ...............................................................................................................................54

III. PLANT PHYSIOLOGY ........................................................................................................................................57

IV. WHAT CUCURBITS ARE GRAFTED .................................................................................................................60

V. GRAFTING METHODS ......................................................................................................................................62

J. Production of Rootstock and Scion ...................................................................................................................67

VI. CONCLUSION .....................................................................................................................................................68

option. Some seed companies now offer watermelon transplants

as 1949, Imazu recommended Cucurbita moschata as a root- TABLE 1

caused by soilborne pests and pathogens when genetic or chem-

less’ fruits have a distinct appearance and a longer shelf life,

Crop Zeatin Dihydo-zeatin Isopentenyl

Wintermelon (Benincasa Lion, Best, Donga Good disease resistance Incompatibility

Methods continue to evolve to overcome existing and devel-

A. Tongue Approach/Approach Graft

Sakata et al., 2007). This method is suitable when rootstock and

cotyledons and the first true leaf start to develop (about 7 to 10

nels inside greenhouses where it is possible to maintain a high patibility toxins.

FIG. 6. Grafting compatibility between scion and rootstock.

57–71. (in Japanese) Fruits and Fertilizer Technology Center PT2004–36.

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