Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide


Cucumber green mottle mosaic virus



Cucumber green mottle mosaic virus


  • Last modified
  • 14 May 2019
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Preferred Scientific Name
  • Cucumber green mottle mosaic virus
  • Taxonomic Tree
  • Domain: Virus
  •   Family: Virgoviridae
  •     Genus: Tobamovirus
  •       Species: Cucumber green mottle mosaic virus
  • Summary of Invasiveness
  • CGMMV is a species of virus in the genus Tobamovirus, which was first described in 1935 in England. Between 1935 and 1985 it spread slowly to other countries, but faster between 1986 and 2006, and rapidly betwe...

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Cucumber green mottle mosaic virus (white break mosaic); symptoms on watermelon plants.
CaptionCucumber green mottle mosaic virus (white break mosaic); symptoms on watermelon plants.
Copyright©Craig Webster/DPIRD, Australia
Cucumber green mottle mosaic virus (white break mosaic); symptoms on watermelon plants.
SymptomsCucumber green mottle mosaic virus (white break mosaic); symptoms on watermelon plants.©Craig Webster/DPIRD, Australia
Cucumber green mottle mosaic virus (white break mosaic); symptoms on cucumber plants.
CaptionCucumber green mottle mosaic virus (white break mosaic); symptoms on cucumber plants.
Copyright©Craig Webster/DPIRD, Australia
Cucumber green mottle mosaic virus (white break mosaic); symptoms on cucumber plants.
SymptomsCucumber green mottle mosaic virus (white break mosaic); symptoms on cucumber plants.©Craig Webster/DPIRD, Australia
Cucumber green mottle mosaic virus (white break mosaic); symptoms on cucumber.
CaptionCucumber green mottle mosaic virus (white break mosaic); symptoms on cucumber.
Copyright©Thorben Lundsgaard/Dept of Plant Biology, KVL, Denmark
Cucumber green mottle mosaic virus (white break mosaic); symptoms on cucumber.
SymptomsCucumber green mottle mosaic virus (white break mosaic); symptoms on cucumber.©Thorben Lundsgaard/Dept of Plant Biology, KVL, Denmark


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Preferred Scientific Name

  • Cucumber green mottle mosaic virus

Other Scientific Names

  • bottlegourd Indian mosaic virus
  • cucumber green mottle mosaic tobamovirus
  • cucumber green mottle mosaic watermelon strain (W)
  • cucumber mottle virus
  • cucumber virus 2
  • cucumber virus 3
  • cucumber virus 4
  • cucumis virus 2
  • tobacco mosaic virus watermelon strain-W

International Common Names

  • English: white break mosaic

Local Common Names

  • Germany: Gurkengrünscheckungsmosaik-Virus

English acronym


EPPO code

  • CGMMV (Cucumber green mottle mosaic tobamovirus)

Summary of Invasiveness

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CGMMV is a species of virus in the genus Tobamovirus, which was first described in 1935 in England. Between 1935 and 1985 it spread slowly to other countries, but faster between 1986 and 2006, and rapidly between 2007 and 2018. It now occurs on all continents except South America. In cucurbits, it causes a damaging disease that reduces fruit yields and quality and spreads efficiently by plant-to-plant contact transmission. Outbreaks occur in many cucurbit crops including vegetables and fruits (e.g. squash and melons). CGMMV seed transmission occurs in at least nine different cucurbit crop species and this is the main way the virus has spread worldwide. Importation of contaminated seeds constitutes a considerable biosecurity concern for counties still without CGMMV. Its high stability and its persistence in contaminated plant material and soil allow it to survive between growing seasons, making eradication difficult.

Taxonomic Tree

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  • Domain: Virus
  •     Family: Virgoviridae
  •         Genus: Tobamovirus
  •             Species: Cucumber green mottle mosaic virus

Notes on Taxonomy and Nomenclature

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Cucumber green mottle mosaic virus (CGMMV) was first described by Ainsworth (1935), although it was previously reported by Bewley (1926) who used the name Cucumis virus 2. CGMMV is a species of the Tobamovirus genus, which is assigned to the Virgoviridae family (Adams et al., 2009). 

Saito et al. (1988) determined the nucleotide sequence of the gene coding for the 30 kDa protein, which is considered to be involved in virus transport between cells. They found homologies with the 30 kDa protein gene of other tobamoviruses. The complete CGMMV amino acid sequence was determined by Meshi et al. (1983), and the amino acid composition by Nozu et al. (1971), Tung and Knight (1972) and Linnasalmi and Toivianinen (1974). The genomic RNA has been sequenced from many isolates (e.g. Ugaki et al., 1991; Kim et al., 2003; Chen et al., 2006).

When partial CGMMV sequences were subjected to phylogenetic analysis, Dombrovsky et al. (2017) reported three major clades. The largest consisted mainly of Chinese CGMMV isolate sequences but also of single isolates from Taiwan, Korea and Japan. The second largest clade consisted solely of Korean CGMMV isolate sequences and the third largest clade consisted of CGMMV sequences of isolates from China, Japan and Taiwan. In addition, a distant minor clade consisted of sequences of isolates from India, Canada and Israel. A second minor, and very distant, clade consisted of CGMMV sequences from Russian and Spanish isolates. The Israeli CGMMV sequence KF155231 from Ecballium elaterium shares a putative ancestor within the Russian/Spanish clade and seemed ancestral to all others.

The responses of Datura stramonium (Solancaeae) and Chenopodium giganteum (Chenopodiaceae) plants to mechanical inoculation were used to distinguish CGMMV strains. The CV3, CV4, Japanese cucumber and watermelon strains all induced local lesions in C. giganteum but failed to infect inoculated leaves of D. stramonium. Although the CV3 and watermelon strains and English C4 failed to infect any Solanaceae species, a German C4 isolate induced chlorotic local lesions in tobacco. The Yodo strain induced local lesions in C. giganteum and D. stramonium, whereas the Indian C strain caused local lesions in C. giganteum. Infected inoculated leaves of D. stramonium were asymptomatic. No systemic infection developed in these indicator host species (Hollings et al., 1975; Dombrovsky et al., 2017 and references therein).


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Infective CGMMV particles are rigid rods ca 300 x 18 nm with a helical structure of pitch 2.3 nm and a central canal of radius 2.0 nm. The central canal is usually clearly visible in the electron microscope in negatively stained preparations. The RNA lies at ca 4.0 nm radius from the centre of the core and the helix comprises 49 subunits/3 turns (Hollings et al., 1975).

The virus has a 6.4 kb single-stranded, positive-sense RNA genome encapsidated within ~2,000 molecules of a single species of capsid protein. The 5′ terminus of the genomic RNA is a methylated nucleotide cap (m7G5′pppG), and a tRNA-like structure is present at its 3′ terminus. These terminal structures protect the ends of their RNA strands from degradation. The genomic RNA contains four open reading frames (ORFs) that encode four defined proteins. Two polypeptides are necessary for its replication complex. First, a 129 kDa polypeptide containing methyltransferase and helicase motifs required for RNA replication. Second, a long 186-kDa polypeptide formed by suppression of a UAG termination codon encodes an RNA-dependent RNA polymerase at its carboxyl terminal domain. Two additional proteins are translated from subgenomic mRNAs corresponding to ORFs in the 3′ half of the genomic RNA. Virion assembly originates in the 3′ terminal ORF encoding the coat protein (CP) (Dombrovsky et al., 2017).


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CGMMV is widespread across Europe and Asia and has also been identified in Nigeria, Canada, USA and Australia.

There is also a record of CGMMV affecting Barbilophozia and Polytrichum mosses and the monocot Deschampsia antarctica from Argentinian Islands in Antarctica (Polischuk et al., 2007). However, considering the unusual hosts and the detection methods used, it must for the moment be considered doubtful. In addition, the record of CGMMV from Brazil (Choudhury and Lin, 1982) has not been confirmed.

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes


ChinaPresentCABI/EPPO, 2015
-GuangdongPresentLi et al., 2009; Liu et al., 2009; CABI/EPPO, 2015
-HebeiPresentLiu et al., 2009; CABI/EPPO, 2015
-HubeiPresentLiu et al., 2009; CABI/EPPO, 2015
-HunanPresentCABI/EPPO, 2015
-JiangsuPresentCABI/EPPO, 2015
-LiaoningPresentZhou et al., 2008; Liu et al., 2009; CABI/EPPO, 2015
-ShandongPresentLiu et al., 2009; CABI/EPPO, 2015
-YunnanPresentCABI/EPPO, 2015
Georgia (Republic of)PresentTarasashvili, 1976; CABI/EPPO, 2015
IndiaPresentNariani et al., 1977; Verma and Verma, 1979; CABI/EPPO, 2015
-DelhiPresentRaychaudhuri and Varma, 1978; CABI/EPPO, 2015
-KarnatakaPresentRashmi et al., 2005; CABI/EPPO, 2015
-MaharashtraPresentCABI/EPPO, 2015
-Tamil NaduPresentCABI/EPPO, 2015; Nagendran et al., 2015
-Uttar PradeshPresentBhargava and Bhargava, 1977; Sandeep and Joshi, 1989; CABI/EPPO, 2015
IranPresentRahimian and Izadpanah, 1977; Rahimian and Izadpanah, 1978; Moradi and Jafarpour, 2011; Nematollahi et al., 2014; CABI/EPPO, 2015
IsraelPresentAntignus et al., 1990; Antignus et al., 2001; CABI/EPPO, 2015
JapanPresentKomuro et al., 1971; Kishi, 1972; Tochihara and Komuro, 1974; CABI/EPPO, 2015
-HokkaidoPresentYoshida et al., 1980; CABI/EPPO, 2015
JordanPresentAl-Tamimi et al., 2009
Korea, Republic ofPresentLee et al., 1990; CABI/EPPO, 2015; Cho et al., 2015
LebanonPresentFaris-Mukhayyish and Makkouk, 1983; CABI/EPPO, 2015
MyanmarPresentKim et al., 2010; CABI/EPPO, 2015
PakistanPresentAli et al., 2004; CABI/EPPO, 2015
Saudi ArabiaPresentAl-Shahwan and Abdalla, 1992; CABI/EPPO, 2015
Sri LankaPresentAriyaratne et al., 2005; CABI/EPPO, 2015
SyriaPresentKassem et al., 2005; CABI/EPPO, 2015
TaiwanPresentWang and Chen, 1985; Chen and Wang, 1986; Hseu et al., 1987; CABI/EPPO, 2015
ThailandPresentCABI/EPPO, 2015
TurkeyPresentGümüs et al., 2004; CABI/EPPO, 2015


NigeriaPresentCABI/EPPO, 2015

North America

CanadaPresent, few occurrencesLing et al., 2014; CABI/EPPO, 2015
-AlbertaPresent, few occurrencesCABI/EPPO, 2015
-OntarioPresent, few occurrencesCABI/EPPO, 2015
USAPresent, few occurrencesCABI/EPPO, 2015
-CaliforniaPresent, few occurrencesNAPPO, 2013; Tian et al., 2014; CABI/EPPO, 2015

South America

BrazilUnconfirmed recordChoudhury and Lin, 1982


AustriaPresentBedlan, 1992; CABI/EPPO, 2015
BulgariaPresentNeshev, 2008; CABI/EPPO, 2015
Czechoslovakia (former)Absent, invalid recordCech et al., 1980; Sindelar et al., 1982; CABI/EPPO, 2015
DenmarkPresentRonde-Kristensen and Jorgensen, 1956; Paludan, 1985; CABI/EPPO, 2015
FinlandPresentLinnasalmi, 1966; Linnasalmi and Toivianinen, 1974; CABI/EPPO, 2015
FrancePresentLetschert et al., 2002
GermanyPresentKohler et al., 1954; Hentschel, 1975; CABI/EPPO, 2015
GreecePresentAvgelis and Vovlas, 1986; Varveri et al., 2002; CABI/EPPO, 2015
-CretePresentCABI/EPPO, 2015
HungaryPresentHorvath, 1992; CABI/EPPO, 2015
LatviaPresentDzirkale et al., 1990; CABI/EPPO, 2015
LithuaniaPresentZitikaite, 2002; CABI/EPPO, 2015
MoldovaPresentCABI/EPPO, 2015
NetherlandsPresentKoot and Dorst, 1959; Dorst, 1988; Runia, 1988; CABI/EPPO, 2015
NorwayPresent, few occurrencesCABI/EPPO, 2015
PolandPresentCABI/EPPO, 2015; Borodynko-Filas et al., 2017
RomaniaPresentPop and Jilaveanu, 1985; CABI/EPPO, 2015
Russian FederationPresentYakovleva, 1965; CABI/EPPO, 2015
-Central RussiaPresentMedvedskaya, 1981; CABI/EPPO, 2015
-Southern RussiaPresentCABI/EPPO, 2015
SpainPresentCélix et al., 1996; CABI/EPPO, 2015
SwedenPresentNilsson, 1977; CABI/EPPO, 2015
UKPresentBewley, 1926; Ainsworth, 1935; Thomas, 1984; CABI/EPPO, 2015
-England and WalesPresentCABI/EPPO, 2015
UkrainePresentKozlov & Poremb'ska, 1972; CABI/EPPO, 2015
Yugoslavia (former)PresentMilicic and Juretic, 1971


AustraliaPresent, few occurrencesCABI/EPPO, 2015; Tesoriero et al., 2016
-Australian Northern TerritoryPresent, few occurrencesCABI/EPPO, 2015; Tesoriero et al., 2016
-QueenslandPresent, few occurrencesCABI/EPPO, 2015
-Western AustraliaPresent, few occurrencesKehoe et al., 2017

History of Introduction and Spread

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Between 1935 and 1985, the known distribution of CGMMV increased within England and spread across Europe to Georgia, Denmark, Germany, Finland, Romania, Russia, Sweden, the Netherlands, Ukraine and former Yugoslavia. It was also found in Lebanon in the Middle East, India in southern Asia, and both Japan and Taiwan in eastern Asia (Dombrovsky et al., 2017 and references therein). CGMMV is presumed to have been introduced to Japan in 1966 by infected Cucurbitaceae seed (Kobayashi, 1990). It was also thought to be introduced through Lagenaria siceraria seed imported from India to Japan (Tochihara and Komuro, 1974). Between 1986 and 2006, CGMMV spread through Europe to Austria, Greece, France, Hungary, Latvia, Lithuania and Spain. In Asia, it spread to Pakistan, Korea and China. During the same period, it was found for the first time in Israel, Saudi Arabia and Syria in the Middle East, and Indonesia and Thailand in South East Asia. During the period 2007 to 2016, CGMMV rapidly dispersed (i) within countries where it was already known, (ii) to more countries in Europe and Asia, and (iii) to other continents. It was reported for the first time in Bulgaria, Moldova, Norway and Poland in Europe, Jordan and Turkey in the Middle East, Iran in central Asia, Sri Lanka in southern Asia, Myanmar in South East Asia, and Nigeria in West Africa. Between 2013 and 2016, it reached Canada and the USA in North America as well as Australia (Dombrovsky et al., 2017 and references therein). This included Alberta and Ontario in Canada (Ling et al., 2014) and California in the USA (Tian et al., 2014). In 2014 to 2017 it spread to three Australian states, the Northern Territory, Queensland, and Western Australia (Tesoriero et al., 2014Dombrovsky et al., 2017).

Risk of Introduction

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Risk Criteria Category

Economic Importance: Moderate
Distribution: Europe, Asia, North America, Africa, Australia
Seedborne Incidence: Moderate
Seed Transmitted: Yes
Seed Treatment: Yes

Overall Risk: Moderate

Notes on Phytosanitary Risk

CGMMV-contaminated seeds are spread via international trade in commercial cucurbit seeds and the international exchange of germplasm. Importation of contaminated seeds poses a biosecurity concern for countries without CGMMV (Dombrovsky et al., 2017 and references therein). The protocol of the International Seed Testing Association involves the testing of 9400 seeds per commercial cucurbit seed sample for CGMMV. With smaller seed lots, 20% of the seeds are tested. Efficient quarantine measures in the form of seed treatment and/or accurate testing should be used.

Early detection and removal of CGMMV-infected cucurbit seedlings, as well as stringent hygiene measures, are critical to avoid introduction of CGMMV to new locations (Dombrovsky et al., 2017 and references therein).

Habitat List

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Terrestrial – ManagedCultivated / agricultural land Secondary/tolerated habitat Harmful (pest or invasive)
Protected agriculture (e.g. glasshouse production) Principal habitat Harmful (pest or invasive)
Disturbed areas Secondary/tolerated habitat Harmful (pest or invasive)
Irrigation channels Secondary/tolerated habitat Harmful (pest or invasive)
Rivers / streams Secondary/tolerated habitat Harmful (pest or invasive)

Hosts/Species Affected

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The majority of plant species infected by CGMMV are in the family Cucurbitaceae. These include both major and minor vegetable and fruit cucurbit species such as rockmelon, cantaloupe and honeydew melon (Cucumis melo), cucumber (Cucumis sativus), watermelon (Citrullus lanatus), zucchini, squash and marrow (Cucurbita pepo), pumpkin (Cucurbita moschata and Cucurbita maxima) and several gourd species (e.g. Benincasa hispida, Lagenaria siceraria, Luffa acutangula, Momordica charantia), which are grown either as crops in their own right or as rootstocks for grafted watermelon (Dombrovsky et al., 2017 and references therein).  At least 15 weed species from different continents have been identified as potential natural CGMMV hosts. These belong to nine different plant families (Dombrovsky et al., 2017 and references therein).

Several cucurbitaceous weeds likely act as reservoir hosts of the virus. Symptomless infection by CGMMV occurs in the weed squirting cucumber (Ecballium elaterium) in Israel, where it acts as an important alternative host of the virus (Antignus et al., 1990). Recently Shargil et al. (2017) reported that infected E. elaterium plants could transmit the virus to melon, cucumber and Nicotiana benthamiana via a bioassay. Horvath (1985b) found the same species to be susceptible following artificial mechanical inoculation, and also reported additional systemically infected cucurbit hosts including: Cyclanthera brachystachya, C. pedata, Melothria pendula (locally), M. scabra, Momordica balsamina, Sicyos angulatus (locally) and Zehneria japonica. Horvath (1985a) subsequently found two further systemically infected hosts, Lagenaria siceraria and Luffa aegyptiaca. However, as Horvath’s studies involved artificial inoculation they do not necessarily reflect true natural hosts. Sandeep and Joshi (1989) reported Momordica charantia as a natural host of the virus. Outside Cucurbitaceae, several natural hosts are known, however they are mostly weeds. For instance, Shargil et al. (2017) found additional asymptomatic hosts including: pigweed (Amaranthus graecizans), A. muricatus, dyer’s cotton (Chrozophora tinctoria), dwarf heliotrope (Moluccella laevis) and ashwagandha (Withania somnifera). Shargil et al. (2017) also demonstrated infection of the seed for pigweed and dwarf heliotrope. Prunus armeniaca was reported as a host of the virus (Cech et al., 1980), but this requires confirmation. Silverleaf nightshade (Solanum elaeagnifolium) and black nightshade (S. nigrum) were reported as hosts but this could not be confirmed. Hovárth (1986) reported systemic infection in Emex australis and E. spinosa following artificial inoculation.

For further information on the natural and experimental host ranges of CGMMV, see Hollings et al. (1975), Shargil et al. (2017) and Dombrovsky et al. (2017).

Growth Stages

Top of page Flowering stage, Fruiting stage, Seedling stage, Vegetative growing stage


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The virus becomes systemic in most infected plants reaching all plant parts, including roots and fruit, and remains able to infect other plants even when symptoms are absent.

In infected foliage and fruit, CGMMV symptoms vary between different cucurbit crop species and cultivars of the same species. In cucumber, green mottling occurs on young leaves and fruit surfaces, and infected plants may collapse. In watermelon, leaf mottling and mosaic develop in young plants, and their stems and peduncles develop brown necrotic lesions. Their foliage may develop a bleached appearance and wilt, and their runners, or even whole plants, may die prematurely. However, foliage symptoms sometimes fade in mature plants, especially in open-field situations. The fruits of infected plants often develop symptoms that render fruit unmarketable, including malformation and internal flesh symptoms of sponginess, rotting and yellowing or dirty red discoloration. In melon, young leaves develop initial mottle and mosaic symptoms that often disappear from mature foliage. Their fruits develop different degrees of malformation, mottling and surface netting. In pumpkin, squash, and zucchini, infected foliage is asymptomatic or leaf mottling and mosaic occur. Pumpkin fruits are always asymptomatic, but squash and zucchini fruits are sometimes externally symptomless and internally discoloured and necrotic. CGMMV symptoms are often indistinct in cucurbit seedlings, except when cotyledons turn yellow. In addition to causing marketable yield losses from poor fruit quality, CGMMV also causes gross yield losses, e.g. 15% and >50% in cucumber and watermelon respectively (Hollings et al., 1975; Dombrovsky et al., 2017 and references therein). Symptomless infection can also induce losses (Kooistra, 1968).

Different CGMMV strains differ in the symptoms they cause. For example, in cucumber, the type strain causes leaf mottling, blistering and distortion and stunted growth. Fruits are usually unmarked, but some strains cause severe fruit mottling and distortion. Some Asian cucumber cultivars show no leaf symptoms but they do suffer yield losses. The aucuba mosaic strain induces bright yellow leaf mottling in cucumbers, with only slight leaf distortion and stunting. Fruits may show yellow or silver-coloured streaks and flecks, especially at higher temperatures. In cucumber, symptoms appear 7-14 days after infection. At low temperatures, when the plants grow more slowly, leaf distortion is more severe (Smith, 1972).

In the field, the virus causes mosaic and occasionally wrinkling, green vein-banding and stunting of muskmelons. In the greenhouse it causes mild chlorosis, mosaic, vein-banding, and at the later stages deformed leaves with blisters (strain CGMMV-M) (Raychaudhuri and Varma, 1978). In Lagenaria siceraria the virus causes mosaic symptoms (VIDE, 1996). In Ecballium elaterium (Antignus, 1990) and other weed hosts (Shargil et al., 2017) the virus causes a symptomless infection.

List of Symptoms/Signs

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SignLife StagesType
Fruit / abnormal shape
Fruit / discoloration
Fruit / lesions: black or brown
Leaves / abnormal forms
Leaves / abnormal patterns
Roots / reduced root system
Stems / necrosis
Whole plant / dwarfing

Biology and Ecology

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Biochemical Properties

Virions contain ca 5% nucleic acid and ca 95% protein. Sub-genomic mRNA is found in virions and in infected cells (VIDE, 1996).

The base composition of the genomes of different CGMMV strains were: strain W: 23.2% G, 24.6% A, 20.6% C and 31.6% U (Okada, 1986); strain CV4: 25.8% G, 25.8% A, 19.3% C and 29.5% U (Okada, 1986); strain CV3: 25.5% G, 25.8% A, 18.3% C and 30.8% U (Okada, 1986).

The infectivity of virions is decreased when deproteinized with proteases, but retained when deproteinized with phenol or detergent. A poly-A region is absent. An additional factor is not required for infectivity. The genome has tRNA-like activity and the genome accepts histidine.

The protein of the virions has a molecular weight of Mr 17,261 and is the coat protein. The method of preparation is described by Fraenkel-Conrat (1957) and Tung and Knight (1972).

Purified preparations sediment as a major infective component with minor components (probably dimers and trimers). Sedimentation coefficient (20,W): 185 S to 195 S. Isoelectric point: approximately pH 4.98. A260/A280: 1.38. Amax(260)/Amin(249): 1.05. Values corrected for light-scattering (Hollings et al., 1975).


Replication does not depend on a helper virus (VIDE, 1996). Two polypeptides are necessary for its replication complex. First, a 129-kDa polypeptide containing methyltransferase and helicase motifs required for RNA replication. Second, a long 186-kDa polypeptide formed by suppression of a UAG termination codon encodes an RNA dependent RNA polymerase at its carboxyl terminal domain. Two additional proteins are translated from subgenomic mRNAs. Virion assembly originates in the 3' terminal ORF encoding the CP (Dombrovsky et al., 2017 and references therein).


Sugimura and Ushiyama (1975) found peripheral vesicles in the mitochondria of protoplasts from infected tobacco cv. Xanthii, 24 hours after inoculation. No other cytopathological effects were noted. Hatta and Ushiyama (1973) made the same observation in cucumber plant tissue cells. Virions in the form of virion-containing crystals are present in the cytoplasm of infected cells. Another cellular change is vesiculation of mitochondria (VIDE, 1996).

Virus-Host Interactions

When cucumber cotyledons that had been inoculated with Tobacco mosaic virus (TMV) were challenge-inoculated with CGMMV, the multiplication of TMV and the size and the number of starch-lesions in the cotyledons remained the same as in single inoculation with TMV. However, inoculation with CGMMV before TMV infection decreased the amount of TMV and the number of starch-lesions (Uekusa et al., 1992). In another study, challenge-inoculation with CGMMV of TMV-infected cucumber cotyledons revealed mechanisms inhibiting cell-to-cell movement of virus and substances, apparently induced in TMV-infected areas (Uekusa et al., 1993).

Complementation of the cell-to-cell spread between different related and unrelated plant viruses was observed by Malyshenko et al. (1989). They found that various tobamoviruses, including CGMMV can facilitate the replication of others in non-permissive hosts, probably by the complementation of transport functions. Results of further studies on transport functions are reported by Mushegian et al. (1989).

Sandeep and Joshi (1989) found that 10-50 days after inoculation with the type strain, there was a decrease in chlorophyll content and chloroplast number and an increase in chlorophyllase activity in Momordica charantia compared with healthy plants. Similar results were found in a study of the same phenomena in Cucumis sativus (Srivastava and Tiwari, 1996).

In soil-grown cucumber the cucumber strain of Tobacco necrosis virus infected the roots of the plant and failed to induce systemic infection unless the plants were already infected by CGMMV (Thomas, 1984).

Sindelar et al. (1982) observed a decrease in weight and an increase of total protein, DNA and RNA in infected cucumber by increasing nitrate content in the nutrient medium, which was also correlated with increasing virus reproduction. The correlation between virus reproduction and nitrate reductase activity was significant.

Virus Stability

All strains of CGMMV are extremely stable. In the sap of Cucumis sativus, the type strain loses infectivity after 10 min at 90°C, the watermelon and Yodo strains lose infectivity at 90-100°C and the Indian strain C loses infectivity at 86-88°C.

The dilution end-point ranges from 1:1,000,000 for the type strain to 1:10,000,000 for the watermelon strain. Infectivity survives for several months at laboratory temperatures and several years at 0°C (Hollings et al., 1975).


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A - Tropical/Megathermal climate Tolerated Average temp. of coolest month > 18°C, > 1500mm precipitation annually
Af - Tropical rainforest climate Tolerated > 60mm precipitation per month
Am - Tropical monsoon climate Tolerated Tropical monsoon climate ( < 60mm precipitation driest month but > (100 - [total annual precipitation(mm}/25]))
As - Tropical savanna climate with dry summer Tolerated < 60mm precipitation driest month (in summer) and < (100 - [total annual precipitation{mm}/25])
Aw - Tropical wet and dry savanna climate Tolerated < 60mm precipitation driest month (in winter) and < (100 - [total annual precipitation{mm}/25])
B - Dry (arid and semi-arid) Tolerated < 860mm precipitation annually
BW - Desert climate Tolerated < 430mm annual precipitation
C - Temperate/Mesothermal climate Tolerated Average temp. of coldest month > 0°C and < 18°C, mean warmest month > 10°C
Cs - Warm temperate climate with dry summer Tolerated Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers
Cw - Warm temperate climate with dry winter Tolerated Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)
Cf - Warm temperate climate, wet all year Tolerated Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year

Latitude/Altitude Ranges

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Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
60 30

Means of Movement and Dispersal

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Natural Dispersal

CGMMV has two main virus dispersion pathways: (i) in protected cropping or in the field and (ii) within seedling nurseries growing cucurbits. CGMMV infects its cucurbit plant hosts systemically i.e. all plant tissues, including foliage, fruits, roots and seeds, contain infectious CGMMV. Its virus particles decompose more slowly in contaminated roots and seeds, so these constitute an important source of infection for future cucurbit crops. Also, symptomless CGMMV-infected weeds, contaminated soils and irrigation water, and nutrient solutions are potential inoculum sources (Dombrovsky et al., 2017 and references therein).

Studies on CGMMV transmission in ebb and flow irrigation systems found that healthy cucurbit crops become infected by virus-contaminated nutrient solutions (Buttner et al., 1995). The virus also occurs in river water used for irrigation due to the presence of virus-contaminated debris in the river (Vani and Varma, 1993). The virus also spreads in protected cropping via surface water used for sprinkle irrigation (Dorst, 1988; Dombrovsky et al., 2017 and references therein).

Other CGMMV sources include infected crop residues (Avgelis et al., 1992). CGMMV was found in manure obtained from cows fed on virus-infected cucurbit fruits (Dorst, 1988). Another study has shown that CGMMV and a number of other plant viruses can pass through the alimentary tract of rodents, such as mice and rabbits, without losing biological activity or particle structure (Kegler et al., 1984; Dombrovsky et al., 2017 and references therein).

Vector Transmission

There are no confirmed reports of specific vector transmission. The virus is not transmitted by the aphids Myzus persicae or Aphis gossypii, nor by cucumber leaf beetle Aulacophora femoralis (Hollings et al., 1975). However, the red pumpkin beetle (Aulacophora fevicollis) is a possible non-specific vector according to detailed studies by Rao and Varma (1984). The regurgitated fluid and excreta of the beetles feeding on infected plants were found to contain infective virions. However, the authors note that the voracious feeding of the beetle means that it is likely that contamination from feeding on infected plants may be lost by the time the beetle finishes eating, and that there is little chance of the virus entering non-lethally injured cells left at the site of feeding. In addition, the honey bee Apis mellifera has also been shown to contribute to the spread of the virus non-specifically via transporting CGMMV from infected plants to healthy plants (Darzi et al., 2018). This was probably due to virions adhering to the bee and entering plants through wounds caused by the bee during pollination. Visualization of pollen using the FISH technique showed no virions adhering directly to the pollen (Shargil et al., 2015).

CGMMV is transmissible experimentally by the parasitic plant dodder, including Cuscuta subinclusa, C. lupuliformis and C. campestris. The aucuba mosaic CGMMV strain is transmitted by C. campestris (Hollings et al., 1975).

Accidental Introduction

Transmission of CGMMV most commonly occurs by mechanical contact between infected plants or contaminated surfaces and healthy plants, for instance through foliage contact, when plants are handled during cultivation, or when infected rootstocks are used in watermelon or cucumber cultivation (Hollings et al., 1975). It can spread readily by contact because its virions are stable and remain infectious on contaminated surfaces for long periods. CGMMV is transmitted readily through tiny wounds that develop when healthy plants brush against infected cucurbit crop plants or weeds, or when they come into contact with virus particle-contaminated plant debris or a wide range of other surfaces, including contaminated machinery, equipment, boxes, tools, hands, clothes and shoes (Dombrovsky et al., 2017 and references therein). If cucurbit seedlings grown from healthy seed stocks are planted in soil contaminated by infective plant debris, the virus can enter through small wounds created in root surface cells and root hairs, thereby infecting the seedling. Other mediators of spread-by-contact transmission include contaminated irrigation water and nutrient solutions, and planting contaminated seeds or transplants. When cucurbit plants are trellised in greenhouses or tunnel houses the virus can spread if string is used to tie them up or knives and sheers used to prune them. In open-field situations, CGMMV can spread to wounded healthy plants when they come into contact with contaminated tractor tyres, agricultural machinery or equipment moving through cucurbit fields.

In nurseries, the spread of CGMMV results from human activities, e.g. grafting and pruning, and by irrigation systems, soil mix, dirt and contaminated surfaces. Healthy cucurbit seedlings become infected after being planted in contaminated soil. When such primary inoculum sources occur, contact transmission can result in rapid secondary spread. Inadvertent distribution of CGMMV-infected seedlings by commercial seedling nurseries and of CGMMV-contaminated seed stocks by commercial seed producers both provide significant virus introduction routes to previously uncontaminated cucurbit (Dombrovsky et al., 2017 and references therein). By harvest, CGMMV can then spread to many crop plant and weed hosts. When this happens, contaminated plant debris remains distributed across the growing area following harvest. Therefore, CGMMV can persist in the absence of growing cucurbit crops via (i) contaminated plant debris buried, scattered on the soil surface, or left in discard piles, any surviving roots from the infected plants and virus particles that remain infectious in the soil and (ii) cucurbit seed contamination, weed host and volunteer cucurbit crop infection and contamination of unsterilized tools, machinery and other surfaces (Dombrovsky et al., 2017 and references therein). CGMMV seed transmission occurs in at least nine species of cucurbits and this is the main way it has become disseminated worldwide through international seed trade and germplasm exchange (Dombrovsky and Smith, 2017).

As root-to-root transmission occurs with other species of tobamoviruses, it is likely to occur with CGMMV even if at a low rate (e.g. Koh et al., 2017). However, transmission due to root contact between infected and uninfected Lagenaria siceraria plants did not take place, but 18% transmission of CGMMV occurred among L. siceraria plants sown in soil mixed with infected plant debris (Rao and Varma, 1984).

When contaminated cucumber and melon seed stocks were directly planted in uninfested soil, transmission only occurred to germinated seedlings occasionally, resulting in a low efficiency of seed transmission (0.1%), but transmission was much higher when contaminated watermelon seeds were sown (1-10%). With cucumber, seed transmission rates >12% were reported. In other investigations that included diverse cucurbit crops, recorded seed transmission rates were 2-3% (Dombrovsky et al., 2017 and references therein).

Seedborne Aspects

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The incidence of CGMMV in cucumber seeds has been reported by various authors at 9.5% (Yakovleva, 1965), 8% (Hollings et al., 1975) and 3-17% (Faris-Mukhayyish and Makkouk, 1983). A 5% incidence was detected in watermelon seeds (Hollings et al., 1975) and CGMMV was detected in the hypocotyls of seedlings grown from seed from diseased watermelon plants (Lee et al., 1990). Komuro et al. (1971) did not detect the virus in watermelon seeds, but did find it in bottle gourd (Lagenaria siceraria) seed.

Using specific CGMMV IgG antibodies and fluorescently labelled secondary antibodies, Reingold et al. (2015) showed that, in infected melon seeds, CGMMV is located in the seed coat and also in the endosperm envelope underlying the seed coat. No signal was detected in the embryo of the seeds, suggesting CGMMV is not present in these tissues. Dombrovsky and Smith (2017) report that tobamovirus particles, including CGMMV, are very stable and infectivity persists for several years.

Effect on Seed Quality

No effect of CGMMV on seed quality has been reported.

Pathogen Transmission

Evidence of seed transmission of CGMMV was reported by Lee et al. (1990) in a study that recovered CGMMV from hypocotyls of seedlings grown from the seed of diseased watermelon plants.

Faris-Mukhayyish and Makkouk (1983) estimated a potential transmission rate of 17% on the basis of ELISA results of 60 individual seeds from infected plants. However, when 100 seeds from infected plants were planted, only 3% of the emerged seedlings actually developed symptoms.

When CGMMV-contaminated cucumber and melon seed stocks were sown directly in virus-free soil, transmission to germinated seedlings only occurred occasionally, resulting in low efficiency of seed transmission (0.1%), but transmission was much higher when contaminated watermelon seeds were sown (1-10%). In one study with cucumber, seed transmission rates of >12% were reported. In other studies involving diverse cucurbit crops 2-3% seed transmission rates were recorded. Seedling-to-seedling transmission can also occur through wounds that develop when infected seedlings brush against healthy ones (Dombrovsky et al., 2017 and references therein).

Seed Treatment

Contamination of seeds by CGMMV is mostly external and so it can be eliminated by dry heat-treatment of seeds for three days at 70°C without impairing seed germination (Hollings et al., 1975). Measures to eliminate endosperm-located CGMMV include immersing seeds in a solution of 15% Na3PO4 for 1 h followed by rinsing in water, or hot air at 70°C for 3 h (Schmelzer and Wolf, 1975). Medvedskaya (1981) recommended hot air at 50-52°C for 3 days followed by 78-80°C for 1 day or, alternatively, soaking in 15% Na3PO4 for 1 h.

In a study in Poland (Macias, 2000), cucumber seeds harvested from CGMMV-infected plants were treated with 1% hydrochloric acid, 0.6% acetic acid, 0.3% sodium hypochlorite, 1% calcium hypochlorite, 1% sodium p-toluenesulfochloramine, 5% calcium oxide, 1% sodium hydrate, 10% sodium phosphate, an experimental seed treatment and seed thermotherapy at 70°C over three days. The heat treatment was efficient with fresh seeds only, whereas two-year-old seeds almost totally lost their ability to germinate. Optimum concentrations for disinfection were: 0.5-1.0% hydrochloric acid, 0.3-0.5% sodium hypochlorite and 10% sodium phosphate.

Seed treatments based on dry heat (e.g. 70°C for 3 days) or disinfectants are used commercially against tobamoviruses in seeds of other plants (e.g. tomatoes) without adverse effects on seed germination. However, although the disinfectants can remove CGMMV from outer seed coats, they cannot remove it from inside the seed (Dombrovsky et al., 2017 and references therein). Work by Reingold et al. (2015) showed that treatment with heat and trisodium phosphate, both commonly used methods, were not completely effective at disinfecting CGMMV in highly infected seed lots.

Seed Health Tests

Seed production crops need to be monitored for CGMMV infection, and samples of harvested seed tested to avoid dispersal of the virus. Importing countries should insist on a representative subsample from every imported seed stock being tested before its release, as in Australia where CGMMV testing of 9400 seedlings/subsample is now mandatory (Dombrovsky et al., 2017 and references therein).


Various ELISA methods have been reported to detect CGMMV in seeds (Clark and Adams, 1977; Kawai et al., 1985; Kobayashi, 1990; Lee and Lee, 1990). The sensitivity of ELISA to detect CGMMV in seed was evaluated by Kawai et al. (1985). An optimized method detected the equivalent of one CGMMV-infected seed in 800 healthy seeds. The authors found testing seed lots of 800 seeds sufficiently accurate for practical purposes. Monoclonal antibodies (MABs) to detect the watermelon strain have been prepared in Japan. Under optimal conditions, the titre of the MABs reached 1:16,000 in ELISA (Takahashi et al., 1989). Lee and Lee (1990) used a protein A-ELISA for detecting CGMMV and other plant viruses. In a comparison of various virus detection procedures (bioassay, Elisa, RT-PCR and HDLPAT), bioassay was reliable only for untreated and heat-treated seeds. For detection of other treated intact seeds, RT-PCR was accurate and sensitive but HDLPAT was the most practical procedure when time and expense were considered (Kim et al., 2000).


CGMMV, Tobacco mosaic virus (TMV), Cucumber mosaic virus (CMV) and Zucchini yellow mosaic virus (ZYMV) were detected from individual fruits and seeds of Capsicum annuum and cucumber by reverse transcription-polymerase chain reaction (RT-PCR) (Choi et al., 1998). The dilution end-points for RT-PCR in crude sap from TMV- and CMV-infected C. annuum leaves and CMV- and CGMMV-infected cucumber leaves were 10-5. However, the amount of PCR product obtained from preparation of ZYMV-infected cucumber leaves was 10-fold lower than those of CMV or CGMMV-infected cucumber leaves. In C. annuum, both TMV and CMV were detected in all parts of the fruit wall tissue, but the yields of PCR products in the fruit stalk and its surrounding tissues were higher than those of the end parts of the fruit. However, in cucumber fruit infected with CMV, CGMMV or ZYMV, the fruit wall tissue and seed located in both stalk and end parts showed higher yields of PCR products than those of intermediate parts. Of five viruses analysed, only TMV was detected in the testa of C. annuum seed, and CGMMV and CMV were detected in the tested cucumber seed.

The sensitivity of ELISA and other tests for the detection of CGMMV in seed was evaluated by Kawai et al. (1985) and Kim et al. (2000).

A protocol for the detection of CGMMV within contaminated seed lots using real-time PCR was published recently (Constable et al., 2017).

Chiang et al. (2012) designed a computer program for estimating the proportion of infected seeds and its confidence limit by group testing in seed health assays, but the procedure has not yet been widely tested.

No conspicuous specific seed symptoms of diagnostic use are reported. Lee et al. (2012) report that, unlike healthy cucumber seeds, infected seeds develop unusual hair-like tissues on seed surfaces, but these are probably of little value for detecting seed infection.

Plant Trade

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Plant parts liable to carry the pest in trade/transportPest stagesBorne internallyBorne externallyVisibility of pest or symptoms
Yes Yes Pest or symptoms usually invisible
Yes Yes Pest or symptoms usually invisible
Yes Yes Pest or symptoms usually invisible
Yes Yes Pest or symptoms usually invisible
Yes Yes Pest or symptoms usually invisible
Yes Yes Pest or symptoms usually invisible
Yes Yes Pest or symptoms usually invisible
Yes Yes Pest or symptoms usually invisible
Yes Yes Pest or symptoms usually invisible

Wood Packaging

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Wood Packaging liable to carry the pest in trade/transportTimber typeUsed as packing
Loose wood packing material Yes
Processed or treated wood Yes
Solid wood packing material with bark Yes
Solid wood packing material without bark Yes

Impact Summary

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Economic/livelihood Negative

Economic Impact

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Of the five cucurbit infecting tobamoviruses, CGMMV is the most economically important and has become a major problem in cucurbit production worldwide. Its impact is primarily through reductions in the yield of marketable fruit with affected fruit sometimes showing sponginess or mottling of the rind. In addition to causing marketable yield losses from poor fruit quality, CGMMV causes gross yield losses of >50% in watermelon. Moreover, yield losses may still occur when infected cucumber foliage is asymptomatic (Dombrovsky et al., 2017 and references therein). Depending on the time of infection of cucumber, yields are reduced by 5-16% and fruit quality due to associated symptoms is considerably reduced (Nilsson 1977). The estimated financial loss for cucumber ranged from 2 to 24%.

Considerable economic losses in watermelon due to infection with the watermelon strain have been reported in Japan. Plants develop severe symptoms and fruit pulp deteriorates (Komuro et al., 1971; Takakuwa et al., 1972; Ugaki et al., 1991).

Outbreaks of the virus are particularly problematic in greenhouse grown cucumbers. For example in Germany, CGMMV affected up to 5% of the crop area in 1974 (Hentschel, 1975), in the Moscow region of Russia it was the most widespread virus (Medvedskaya, 1981), in Georgia its incidence in cucumber was 80-100% in some districts (Tarasashvili, 1976) and in Latvia it caused one of the most common virus diseases (Dzirkale et al., 1990). The incidence in some districts of northern India was up to 100% in Lagenaria siceraria, 80% in muskmelon and 75% in watermelon crops by the end of the growing season (Vani and Varma, 1993). In Israel since 2007, CGMMV has become the dominant tobamovirus infecting greenhouse grown trellised cucumbers, causing large scale losses. This rapid rise in incidence has supplanted the previously common cucumber fruit mottle mosaic virus, to the point where only CGMMV is now found (Reingold et al., 2013, 2015, 2016).

Environmental Impact

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No environmental impacts have been recorded but populations of wild cucurbitaceous plant species are likely to become infected when grown in close proximity to cucurbit crops. They can also act as reservoirs of infection for spread to cucurbit crops.

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
  • Benefits from human association (i.e. it is a human commensal)
  • Has high reproductive potential
  • Reproduces asexually
Impact outcomes
  • Host damage
  • Negatively impacts agriculture
  • Negatively impacts livelihoods
  • Negatively impacts trade/international relations
Impact mechanisms
  • Pathogenic
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally
  • Difficult to identify/detect as a commodity contaminant
  • Difficult to identify/detect in the field
  • Difficult/costly to control


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It is seldom possible to identify CGMMV with certainty from symptoms in leaves and fruit of cucurbitaceous species or by indicator plant reactions, as similar symptoms can be caused by other cucurbit infecting viruses. CGMMV-infected plants can be identified in a variety of ways. The most widely used of these methods is based on the polymerase chain reaction (PCR, or derivatives of the technique) including real-time PCR based methods, or antibody based methods such as ELISA. The advantage of PCR-based methods is their high sensitivity and specific detection of the virus (e.g. Ugaki et al., 1991; Yoon et al., 2008; Liu et al., 2009; Kim et al., 2010; Moradi and Jafarpour, 2011; Lee et al., 2012; Beredson and Osterhof, 2015; Reingold et al., 2015) and they are commonly used when high sensitivity is required, such as seed testing programs to ensure freedom from the virus. Antisera specific for the virus are also readily available commercially and also offer later flow device tests for the virus, which provide rapid detection albeit at a lower sensitivity than traditional plate format ELISA. Monoclonal antibodies have also been developed for the virus (Shang et al., 2011) and utilized in a number of formats, with immunocapture RT-PCR (IC- RT-PCR) being up to 400 times more sensitive than traditional double antibody sandwich ELISA or dot immuno-binding assays. Kan et al. (2010) reported that Immunomagnetic Separation (IMS) followed by RT-PCR is a very sensitive method of detecting CGMMV in infected seeds, with an eightfold increase in sensitivity compared to IC-RT-PCR, though it is not yet widely used. 

Li et al. (2013) also reported a reverse transcription loop-mediated isothermal amplification (RT-LAMP) based test for the virus, which offers highly sensitive detection of the virus (100-fold higher sensitivity than RT-PCR). LAMP format assays are highly sensitive and rapid, require minimal equipment, and are ideal for use in field and quarantine situations, but have yet to be implemented widely.

Similarities to Other Species/Conditions

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Viruses with serologically related virions to CGMMV include: Tobacco mosaic virus (TMV); Tomato mosaic virus; Kyuri green mottle mosaic virus (KGMMV), Frangipani mosaic virus, Ribgrass mosaic virus and Tobacco mild green mosaic virus (VIDE, 1996). Viruses with serologically unrelated virions include: Odontoglossum ringspot virus and Sunn-hemp mosaic virus (SHMV) (VIDE, 1996). Ullucus mild mottle virus, a distinct Tobamovirus, is closely related to some tobamoviruses, including CGMMV (Offei et al., 1995). 

A comparison of 126 kDa proteins for Tobamovirus classification was made by Fraile and Garcia-Arenal (1990). The 126 kDa protein, which is considered to be a part of the viral replicase, was translated in vitro, and a comparison was made between eight tobamoviruses by SDS-PAGE. It grouped CGMMV together with SHMV and KGMMV.

In determining the complete nucleotide sequence of the SH strain of CGMMV, Ugaki et al. (1991) found that the sequence showed 55-56% identity to that of strains of TMV and Tobacco mild green mosaic virus.

The host ranges of CGMMV and KGMMV are similar. Their coat proteins have greater amino acid sequence similarity than either virus has to other tobamoviruses, which are serologically more distantly related (VIDE, 1996).

In more recent phylogenetic studies on members of the Tobamovirus genus, the five cucurbit-infecting viruses within the Cucurbitaceae-infecting group formed a major phylogrouping on their own. KGMMV, Zucchini green mottle mosaic virus (ZGMMV) and Cucumber fruit mottle mosaic virus (CFMMV) all grouped together. In contrast, CGMMV and Cucumber mottle virus (CMoV) each formed their own monophyletic minor groups. Research showing the separation of CGMMV as a monophyletic group agreed with earlier research that it was a divergent Tobamovirus species (Gibbs et al., 2015; Dombrovsky et al., 2017 and references therein).

Prevention and Control

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Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

Integrated disease management (IDM) approaches that combine phytosanitary (hygiene), cultural (agronomic), chemical (against vectors), host resistance and biological control measures that are most appropriate for a given virus-host pathosystem and growing situation, provide the best means of plant virus disease control. Effectiveness is optimized by including measures that are complementary and act in as many different ways as possible to target vulnerable components of the disease cycle. This includes deploying measures with low and high selectivity that address external and internal virus sources as well as early and late virus spread. However, the ability to combine control measures is limited by cost, and affordability is greatest with high value protective cropping and lowest in low value open-field situations. With CGMMV, IDM development is simplified by the lack of a specific invertebrate or fungal vector. It relies primarily on diverse phytosanitary control measures, but cultural, biological and host resistance measures are also available (Dombrovsky et al., 2017 and references therein).

Phytosanitary Measures

To avoid its introduction, virus-free seed sources are required. However, once it is present in a location, additional measures are necessary to limit its spread. Seed treatments with sodium triphosphate or heat greatly reduce the level of infection, but are not sufficient to completely eliminate the virus in highly contaminated seed lots (Reingold et al., 2015). The high stability of the virus enables its survival on surfaces, as well as within soil from infested properties. Therefore, reducing levels of CGMMV contamination and the degree of contact of plants with infectious material are both essential (Reingold et al., 2016).

To avoid CGMMV spread once it becomes established on a farm, all contaminated disposable materials (such as plastic mulch, wooden stakes and string) and organic materials (such as crop debris, weeds and cull piles) should be destroyed by incineration or deep burial. Virus-contaminated agricultural infrastructure, such as metal stakes, posts, containers, equipment and machinery, should be subjected to high-pressure cleaning with a detergent/disinfectant to ensure that they are free of soil and plant material. Disinfectant should be used to clean any irrigation lines, pipes and water tanks. Sites chosen for cleaning must be located where waste water and debris can be safely disposed of by running it into a covered drain or hole that avoids contamination of the water table. Once they have been disinfected, cleaned items should be stored away from any possible sources of recontamination. Unless disinfected, agricultural machinery and equipment should never be moved from infected to healthy crops or between contaminated and uncontaminated farms or protected-cropping facilities. To prevent contamination from visiting trucks, other vehicles and people, access should be limited to a secure place near the entrance to a farm and where cleaning and disinfection facilities are present. Eliminating any potential CGMMV reservoirs in alternative host weed and volunteer cucurbit crop plants is crucial to prevent carryover between successive cucurbit growing seasons. Methyl bromide fumigation removes the virus from infected soil but is no longer recommended because of the damage it can cause to the environment. Using stabilized chlorine with the active ingredient troclosene sodium (C3Cl3N3NaO3) to destroy CGMMV is important when transplanting cucurbit seedlings into CGMMV-contaminated land and when recycling water before reuse (Dombrovsky et al., 2017 and references therein).

In seedling nurseries, early detection and removal of CGMMV-infected cucurbit seedlings, as well as stringent hygiene measures, are critical to avoid its introduction to new farms. CGMMV spread by people undertaking grafting, or other procedures requiring plant handling, must be avoided by using disposable gloves and regular cleaning of hands and clothing using detergent sanitizers. Robots reduce the cost of grafting onto cucurbit rootstocks and are easy to disinfect. Another example of  a newer technology that offers exciting prospects for phytosanitary control is remote sensing via lightweight unmanned aerial vehicles and precision agriculture (i) on a micro-scale to identify and remove infected cucurbit seedlings before their distribution from nurseries to end-users or (ii) on a macro-scale to provide valuable information about virus infection foci within individual crops and to know where to target control measures (Dombrovsky et al., 2017 and references therein).

A variety of chemical agents are reported to be effective in treating Tobamovirus contamination. Oxidizing agents like bleach are highly effective; however their use on certain surfaces and in certain environments is problematic due to their harsh effect on metal surfaces and their risk to skin, eyes etc. Other popular treatments include commercially available disinfectant solutions, non-fat milk powder and trisodium phosphate, which are effective for a variety of viruses and viroids including tobamoviruses (e.g. Lewandowski et al., 2010; Li et al., 2015). Steam heat or ozone treatments and ultraviolet irradiation can also be used to destroy infectious CGMMV. However, there is an urgent need to diversify the range of currently available disinfectants/virucidal chemicals effective against this virus (Dombrovsky et al., 2017 and references therein).

Ozonisation, ultrafiltration and heat treatment of drain water from soilless cultures were tested to assess their efficacy for the elimination of pathogens, including CGMMV. Ultrafiltration and heat treatment at more than 90°C gave promising results, allowing the drain water to be reused without risk (Runia, 1988). By lowering the pH in the water to pH 4 and by using a better ozone distribution, CGMMV was completely eliminated in 75 min with 6 g ozone/h (Runia, 1988). Ozone installation was used by many growers in the Netherlands in 1994 (Runia, 1994).

Vani and Varma (1993) recommend the burning of plant debris to avoid transmission from CGMMV contaminated material to new plantings. Composting of infected cucumber plants resulted in elimination of the virus. The temperature of 72°C reached during the composting was thought to be the main reason for the elimination (Avgelis et al., 1992). Disinfection of hands and utensils by a solution of soft soap and trisodium phosphate (Na3PO4) (200 g and 50 g, respectively, in 1 L of water) was recommended by Schmelzer and Wolf (1975).

Cultural Measures

All field, tunnel house or greenhouse sites where CGMMV-infected cucurbits have been present must be considered contaminated. A minimum of 24 continuous months should pass before any cucurbits are grown again. Non-host crops should be planted instead, preferably ones that manage soil erosion and water run-off, such as millet, maize or sorghum in tropical or subtropical regions, and thickly sown barley, oats, grass pasture or alfalfa in temperate regions. Weed and volunteer cucurbit control should be rigorous to prevent any alternative host CGMMV carryover (Dombrovsky et al., 2017 and references therein).

Host-Plant Resistance

Both complete and partial CGMMV resistance occur in cucumber. Recessive polygenes confer partial resistance in melon. In addition, strain-specific recessive resistance genes cgmmv-1 and cgmmv-2 in melon accession Chang Bougi, all attenuate CGMMV symptoms. CGMMV resistance also occurs in Cucumis africanus, C. ficifolius, C. meeusei, C. myriocarpus and C. zeyheri (Dombrovsky et al., 2017 and references therein).

In wild Cucumis species, absolute resistance was found in C. ficifolius. C. melo var. momordica was also resistant (Pan and More, 1996). Rajamony et al. (1990a) studied inheritance of resistance in C. melo. Rajamony et al. (1990b) screened 187 collections of C. melo and eight wild Cucumis species, but only a few melon species were found to be resistant. African Cucumis species were studied for resistance by Visser (1986). In testing 11 wild cucurbit species for resistance, C. africanus was completely resistant (Kiessling et al., 1985). Inheritance of resistance in C. anguria was investigated by Nijs (1982). When screening a range of Cucurbita accessions, Horvath (1992) found immunity in 24 out of 25 accessions of C. moschata. Hsiao et al. (1993) summarized 10 years of screening work on cucurbit crops (174 cultivars) in Taiwan for resistance to viruses, including CGMMV. Cucumis ficifolius was resistant in an evaluation of C. melo and wild relatives (Osaki et al., 1994). Progressive stages and results of breeding work for resistance in melon are described by More et al. (1993). The introduction of resistance in cucumber was reported by Teploukhova (1986).

Grafting cucurbit scions onto CGMMV-resistant rootstocks, consisting of bottle gourd, various Cucurbita species and interspecific crosses between pumpkins and resistant melons, is a method used commercially to prevent soilborne CGMMV infections. For example, in 2016, to prevent such infections, Hishtil nurseries in Israel sold 1.5 million watermelon seedlings grafted onto CGMMV-resistant rootstocks (Dombrovsky et al., 2017 and references therein).

Apart from natural resistance, other forms of resistance offer prospects for CGMMV control in the future. One involves introducing small interfering RNA directly into plants. Another involves developing transgenic plants that overexpress virus CP genes (Dombrovsky et al., 2017 and references therein). Emran et al. (2012) have produced resistant melon plants by transferring direct repeats of the CGMMV coat protein gene into susceptible plants. Plants containing the transgene were highly resistant to infection due to RNA silencing. A similar approach involves grafting commercial cucurbit cultivars onto watermelon rootstocks with transgenic CGMMV resistance. However, a drawback to this is that when the grafted transgenic rootstock is exposed to infection, CGMMV is transferred from wounded roots to the soil. When watermelon rootstocks were transformed with viral CP using Agrobacterium, CGMMV resistance was conferred to fruits. CGMMV CP mRNA and protein was expressed in the transgenic watermelon rootstock but not in the watermelon scions. The clustered regulatory interspaced short palindromic repeat (CRISPR)-Cas9 system provides a different approach to achieving future CGMMV resistance that has been adapted to interfere with protein synthesis from RNA viruses like CGMMV (Dombrovsky et al., 2017 and references therein).

Biological Measures

Possible protection by pre-inoculation with the slightly pathogenic strain S7 of Tobacco mosaic virus (TMV) onto cucumber was examined by Vlasov and Parshin (1981). A detailed study is reported on the isolation of an attenuated form (SH33b) of CGMMV. Its properties to protect muskmelon under experimental and cultivation conditions were investigated. The properties were compared with those of an attenuated strain of TMV used to cross-protect tomato (Motoyoshi and Nishiguchi, 1988). This approach is currently being developed for commercial CGMMV control in Japan and Russia (Dombrovsky et al., 2017 and references therein).


An inhibitor (a protein) found in extracts of Mirabilis jalapa showed highly potent activity against the mechanical transmission of CGMMV (Kubo et al., 1990). Hippuric acid altered the host susceptibility towards the virus according to a detailed study by Verma and Verma (1979). Leaf extracts of Pseuderanthemum bicolor also showed an inhibitory effect against CGMMV (Verma and Khan, 1984). Tannins isolated from the bark of several trees were studied and found to have enzyme-inhibitory and anti-plant-viral activities (Zhang et al., 1990). Of several substances sprayed on test plants, acetone extracts of Phytolacca acinosa were the most effective in inhibiting infection of CGMMV (Kuwata et al., 1979). The application of an airbrush-inoculation technique for screening anti-viral substances against CGMMV is described by Huang et al. (1974). An extract from dried roots of mature plants of Boerhavia diffusa or a glycoprotein isolated from the roots prevented infection of melon when sprayed on leaves before inoculation (Awasthi et al., 1984).

Gaps in Knowledge/Research Needs

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The eight future research needs suggested by Dombrovsky et al. (2017) are as follows:

  1. To avoid further inadvertent dispersal to new countries via international seed trade, crops grown for cucurbit seed production should be monitored for CGMMV and representative subsamples of seed tested for it. Importing countries need to ensure that a representative subsample from every imported seed stock is tested before release by a suitably accredited laboratory.
  2. Thorough application of phytosanitary measures is critically important for seedling nurseries as dispersion via infected cucurbit seedlings constitutes a key risk pathway for CGMMV contamination of farms and protected cropping facilities.
  3. There is a need to diversify the range of currently available disinfectants/virucidal chemicals effective against CGMMV.
  4. Research is required on how long CGMMV infectivity persists in (i) soils as free virus particles or inside plant debris and live roots and (ii) irrigation water and nutrient solutions. 
  5. More research is required to establish whether wind-assisted contact transmission occurs in cucurbit crops and if additional beneficial insects, other than honey bees, are responsible for spreading CGMMV via pollination or plant wounding.
  6. More information is needed about the extent CGMMV survives outside the cucurbit growing season in alternative weed host species in different world regions, and whether carry over occurs via seed-borne infection in weeds.
  7. Improved routine procedures for large-scale CGMMV detection in seed, seedling, soil and water samples will help ensure its rapid and accurate diagnosis.
  8. Newer technologies offer prospects for greatly improved CGMMV management, e.g. (i) remote sensing via lightweight unmanned aerial vehicles and precision agriculture on a micro-scale to identify and destroy infected cucurbit seedlings in nurseries and a larger scale to identify where to target phytosanitary control measures in large-scale production, (ii) exploiting CRISPR-Cas9 methodology to provide an alternative approach to conventional breeding for CGMMV resistance in new cucurbit cultivars or (iii) employing robots to reduce the cost of grafting onto cucurbit rootstocks in nurseries.


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22/05/18 Updated by:

Craig Webster, Department of Primary Industries and Regional Development, Agriculture and Food, Western Australia, Australia

Roger Jones, Department of Primary Industries and Regional Development, Agriculture and Food, Western Australia, Australia

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