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(Upload on September 11 2016) [ 日本語 | English ]

1892 Iwanovsky WP (USSR): discovering that tobacco mosaic virus (TMV) was filterable 1898 Beijerinck: porposed virus (L.) 1935 Stanley M 1904-1971, USA TMV = globulin-like protein (corrected later → protein + nucleic acid) exstracted and crystallized the virus that was acicular-crystallied nucleic protein wiht 40 million MW, by using enzymatic chemistry 1936 Sir F Bowden & Pirie: confirmed that TMV was nucleoprotein 1939 Pfankuck et al.: emphasized the importance of morphology (phenotypic classification) after the detailed observation of TMV 1953, 1957 Lwoff: summarized the characteristics of virus 1) present either DNA or RNA 2) replicable by nucleic acids 3) not grown and not reproduced by binary division 4) absent energy-supply system 5) utilizing the ribosomes of host (obligate parasitic) Virus groups based on the hosts Bacterial virus invertebrate virus Vertebrate virus Plant virus

[classification, structure, biology] Pathway of bacteriophage T4 morphogenesis. Genes participating in assembly are numbered. Chaperones that are not part of the final structure are presented in parentheses.

The families of virus infecting bacteria Non-enveloped_____________________________________| Enveloped The families of virus infecting plants Non-enveloped_____________| Enveloped

The families of virus infecting invertebrates Non-enveloped_____________| Enveloped The families of virus infecting vertebrates Non-enveloped_____________| Enveloped

Taxonomy (分類)

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Table II. List of virus families and groups in order of presentation (proposed in 1980). V: vertebrates, I: invertebrates, P: plants. ds: double strand, ss: single strand. *: some members possess a cell membrane-derived envelope.

Characterization		Families or groups	Kinds of host
dsDNA, enveloped		Poxviridae Herpesviridae Baculoviridae Plasmaviridae	V, I V I bacteria
dsDNA, non-enveloped	tailed phages	Iridoviridae * Adenoviridae Papovaviridae Caulimovirus Tectiviridae Corticoviridae Myoviridae Styloviridae Podoviridae	V, I V V P bacteria bacteria bacteria bacteria bacteria
ssDNA, non-enveloped		Parvoviridae Geminivirus Microviridae Inoviridae	V, I P bacteria bacteria
dsRNA, enveloped		Cystoviridae	bacteria
dsRNA, non-enveloped	possible family possible family possible family possible family	Reoviridae Isometric dsRNA mycoviruses requiring only one RNA segment for replication Isometric dsRNA mycoviruses requiring two RNA segments for replication Isometric dsRNA mycoviruses requiring three RNA segments for replication Bisegmented dsRNA viruses	V, I, P fungi fungi fungi V, I
ssRNA, enveloped Genome strategy	a. No DNA step (i) positive-sense genome (ii) negative-sense genome (iii) not established b. DNA step in replication cycle Bipartite genomes Isometric particles Possible group Rod-shaped particles Tripartite genomes Isometric particles Isometric and bacilliform particles Rod-shaped particles	Togaviridae Coronaviridae Paramyxoviridae Orthomyxoviridae Rhabdoviridae Bunyaviridae Arenaviridae Tomato spotted wilt virus group Retroviridae Dianthovirus Comovirus Nepovirus Pea enation mosaic virus group Nodaviridae Velvet tobacco mottle virus group Tobravirus Cucumovirus Bromovirus Ilarvirus Alfalfa mosaic virus group Hordeivirus	V, I V V V V, I, P V, I V P V P P P P I P P P P P P P
ssRNA, non-enveloped	Monopartite genomes Isometric particles Rod-shaped particles	Picornaviridae Caliciviridae Nudaurelia β virus group Leviviridae Maize chlorotic dwarf virus group Tymovirus Luteovirus Tombusvirus Sobemovirus Tobacco necrosis virus group Closterovirus Carlavirus Potyvirus Potexvirus Tobamovirus	PV, I V I bacteria P P P P P P P P P P P

ICTV (International Committee on Taxonomy of Viruses) (2015)

Ord Caudovirales

Fam Myoviridae

Fam Podoviridae

Fam Siphoviridae

Ord Herpesvirales

Fam Alloherpesviridae

Fam Herpesviridae

Fam Malacoherpesviridae

Ord Ligamenvirales

Fam Lipothrixviridae

Fam Rudiviridae

Ord Mononegavirales

Fam Bornaviridae

Fam Filoviridae

Fam Mymonaviridae

Fam Nyamiviridae

Fam Paramyxoviridae

Fam Pneumoviridae

Fam Rhabdoviridae

Fam Sunviridae

Fam Unassigned

Ord Nidovirales

Fam Arteriviridae

Fam Coronaviridae

Subfam Letovirinae: Alphaletovirus Subfam Orthocoronavirinae (= Coronavirinae, coronavirus): Alphacoronavirus, Betacoronavirus, Deltacoronavirus, Gammacoronavirus

Fam Mesoniviridae

Fam Roniviridae

Ord Picornavirales

Fam Dicistroviridae

Fam Iflaviridae

Fam Marnaviridae

Fam Picornaviridae

Fam Secoviridae

Fam Unassigned

Ord Tymovirales

Fam Alphaflexiviridae

Fam Betaflexiviridae

Fam Gammaflexiviridae

Fam Tymoviridae

Virus families not assigned to an Ord Fam Adenoviridae Fam Alphatetraviridae Fam Alvernaviridae Fam Amalgaviridae Fam Ampullaviridae Fam Anelloviridae Fam Arenaviridae Fam Ascoviridae Fam Asfarviridae Fam Astroviridae Fam Avsunviroidae Fam Baculoviridae Fam Barnaviridae Fam Benyviridae Fam Bicaudaviridae Fam Bidnaviridae Fam Birnaviridae Fam Bromoviridae Fam Bunyaviridae Hantaviruses: single-stranded, enveloped, negative sense RNA viruses that can kill humans Fam Caliciviridae Fam Carmotetraviridae Fam Caulimoviridae Fam Chrysoviridae Fam Circoviridae Fam Clavaviridae

Fam Closteroviridae Fam Corticoviridae Fam Cystoviridae Fam Endornaviridae Fam Flaviviridae Fam Fuselloviridae Fam Geminiviridae Fam Genomoviridae Fam Globuloviridae Fam Guttaviridae Fam Hepadnaviridae Fam Hepeviridae Fam Hypoviridae Fam Hytrosaviridae Fam Inoviridae Fam Iridoviridae Fam Lavidaviridae Fam Leviviridae Fam Luteoviridae Fam Marseilleviridae Fam Megabirnaviridae Fam Metaviridae Fam Microviridae Fam Mimiviridae Fam Nanoviridae Fam Narnaviridae Fam Nimaviridae Fam Nodaviridae Fam Nudiviridae

Fam Ophioviridae Fam Orthomyxoviridae Fam Papillomaviridae Fam Partitiviridae Fam Parvoviridae Fam Permutotetraviridae Fam Phycodnaviridae Fam Picobirnaviridae Fam Plasmaviridae Fam Pleolipoviridae Fam Polydnaviridae Fam Polyomaviridae Fam Pospiviroidae Fam Potyviridae Fam Poxviridae Fam Pseudoviridae Fam Quadriviridae Fam Reoviridae Fam Retroviridae Fam Sarthroviridae Fam Sphaerolipoviridae Fam Spiraviridae Fam Tectiviridae Fam Togaviridae Fam Tombusviridae Fam Totiviridae Fam Turriviridae Fam Unassigned Fam Virgaviridae ICTV (2015)

Structure (構造)

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TMV (タバコモザイクウィルス) 300 nm in height × 18 nm in width rod-shaped virus → applicable this basic structure to rod-shaped and string-shaped viruses present 2130-17500 dt of proteins components: protein = 95% + RNA 5% Fig. Structure of TMV. (A) Drawing to show the relationships of the RNA and the protein subunits. The DNA shown free of protein could not maintain this configuration in the absence of the protein subunits. There arc 3 nucleotides per protein subunit, or 49 per turn of the major helix spaced about 5 A apart. (B) Photograph of a model of TMV with major dimensions indicated. The structure repeats after 69 A in the axial direction, the repeat containing 49 subunits distributed over 3 turns of the helix.(Klug & Caspar 1960, and based largely on the work of R. E. Franklin) (C) Photograph of a particle of TYMV obtained by symmetrizing an electron microgrpah of a single particle. (Markham 1968) The molecular weights of basic proteins differ between this TMV and the other TMV. Many species have proteins of which molecular weights are approximately 18-Kd → attempting to apply virus classification

Rhabdovirus Enveloped, bullet shaped. 180 nm long and 75 nm wide. Certain plant rhabdoviruses are bacilliform in shape and almost twice the length. The basic structure is comparable with the structure of TMV. → Viron

Spherical virus (球形ウィルス) All spherical virus shells have Icosahedral symmetry = Built from 20 identical equilateral triangles. → enclose maximum volume → probably structurally most stable The structures were predicted by of subunit sequences The surface structures of spherical virus is estimated by the variations of the combinations of subunits and/or the relationships between variations of subunit shapes and numbers of subunits Fig. A. Comparison of models each built of 180 units (corks) on the surface of a sphere representing in position, direction and radial extent the structure units in the outer regions of thc protein shells of A BBMV, B TYMV (Finch & King 1966). 1) Twofold view, 2) threefold view, 3) fivefold view. The structure units of BBMV protrude ca 15 A from the body of the particle and lie with their long axes approximately parallel to the nearest local sixfold or fivefold axis of the particle, forming 32 cylindrical or prismatic morphological units. The structure units of TYMV protrude further from the body of the particle than do those of BBMV and are tilted further toward the nearest local sixfold or fivefold axis to form morphological tin its shaped like frusta of cones. (Finch & King 1967) Fig. B. Clustering patterns of subunits in icosahedra. Top left is a simple icosahedron. Its triangular faces are undivided (T = 1): each is defined by three subunits and thus there are 60 subunits in the entire icosahedron. These subunits may cluster in fives (a, pentamer clusters) at the vertices as in the particles of satellite virus. Middle left is an icosadeltahedron of 180 subunits. Each triangular face of the underlying icosahedron (broken lines) is divided into six half triangles (unbroken lines: T = 6 × 1/2 = 3). Each triangular face is defined by three subunits, and these could cluster together (trimers), or may cluster with subunits of adjacent triangles either in pairs (dimers), or in fives and sixes (pentamer/hexamers). Dimer clustering gives, in one view, a 'Star of David' pattern (b), and this is found in particles of turnip crinkle virus. Pentamer/hexamer clustering (d) is seen in particles of tymoviruses. The particles of some viruses, such as alfalfa mosaic virus are tubular variants of an icosadeltahedron (c). These particles consist of two half icosadeltahedra joined by a tubular net, which has no five-fold axes. The clustering pattern in alfalfa mosaic virus particles is not known for sure; the subunits may be clustered as pentamer/hexamers (c. top half) or trimers (c. lower half).

A B

Plant virus (植物ウィルス)

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Multicomponent and segmental genome Only seen in plant virus Information expression mechanisms on genome or gene genome (= nucleic acids) → enzyme, protein (coat protein) = form, etc. TMV = 5% nucleic acids + 95% proteins TYMV = 34% nucleic acids TNV satellite virus = 20% nucleic acids – cannot reproduce without TNV PVX = 6% nucleic acids + 94% proteins PVY = 5% nucleic acids

Table 1. Arrangement of elongated plant viruses accroding to normal length (NL), shape, diameter (φ), mode of transmission, and thermal inactivation point (TIP).

Group	NL (rnp)	Shape	φ (μm)	Transmission	TIP- (°C)	a	b
1	130	Rigid rods	20	Sap; soil	-	Barley stripe mosaic virus (1)	Soil-borne wheat mosaic virus (2)
2	180	Rigid rods			75-80	Potato stem mottle virus (3)
3	300	Rigid rods	15	Sap	85-95	+ Tobacco mosaic virus (4) + Cowpea virus (5)	+ Cucumber green mottle virus (6) Odonioglossum ring spot virus (7)
4	480	Flexible threads	10-11	Sap; (aphids?)	60-75	White clover mosaic virus (8)	Cymbidium mosaic virus (7)
5	515	Flexible threads				Potato virus X (9)	Cactus virus 1 (10)
6	580	Flexible threads				Potato aucuba mosaic virus (11)
7	620	Rods, rigid to slightly flexible	12-13	Apids; sap	65-75	Wisconsin pea stieak virus (12)
8	650	Rods, rigid to slightly flexible				Red clover vein mosaic virus (12) * Carnation latent virus (13) * Potato virus S (13) * Potato virus M (13) Cactus virus 2 (10)	"Hop virus" (14) "Gladiolus virus"(14) "Poplar virus" (14) Tulip-breaking virus (15)
9	700	Rods, rigid to slightly flexible	12-13	Mites; sap	-	Wheat streak mosaic virus (16)
10	730	Flexible threads	12-13	Aphids; sap	50-60	Beet mosaic virus (17) Potato virus A (18) Potato virus Y (18) Tobacco etch virus (19) Henbane mosaic virus (19)
11	750	Flexible threads				‡ Bean common mosaic virus (20) ‡ Bean yellow mosaic virus (20) ‡ Pea mosaic virus (20) Soybean niosaic virus (20) Turnip mosaic virus (21) Cockaf not streak virus (19) Lettuce aulusaic virus (22) Sorghum red stripe virus (23)	Sugar cane mosaic virus (24)
12	1250	Very flexible threads	10	Aphids; (sap)	-	Beet yellows virus (25)

a, b, +, *, ‡ Indicate serological relationship within groups 3, 8, and 11, respectively. Numbers in parentheses refer to footnotes as follows: (1) Gold et al., 1954: 130 × 30 μm. Brandes (1959) gave a NL of 128 μm, but the diameter of the particles was only about 20 μm when compared with TMV and potato stem mottle virus. (2) Gold et al., 1957. (3) Paul & Bode (1955) found two frequent lengths: 70 and 180 μm. The values of Harrison & Nixon (pers comm) are in good accordance. These authors could show that the shorter particles are noninfectious. Therefore, 180 μm must be taken as NL. Potato stem mottle virus also includes tobacco rattle virus and soil-borne potato viruses found in the United States (Walkinshaw & Larson 1958, Oswald & Bowman 1958). The morphological data are apparently the same; furthermore, serological relationship has been established. (4) A great number of papers exist; 300 μm is especially the result of detailed measurements by Williams & Steers (1951). TMV often has been used as a standard in our laboratory. (5) The cowpea virus was kindly supplied by FC Bawden. (6) A detailed study on this subject is that of Knight (1955). (7) Jensen & Gold, 1951; Newton & Rosberg, 1952; Gold & Jensen, 1953; Murakishi, 1958. (8) Brandes & Quanta, 1957. (9) Bode & Paul, 1955. (10) With the aid of the dipping method we found two different viruses in several species of cactus plants (Brandes & Uachdraweit, unpublished). Cactus virus 1 possibly is identical with a virus described by Amnelunxen (1958). (11) Paul & Bode, 1956. (12) Wetter et al., 1959; Wetter & Quanta, 1953; Brandes & Quanta, 1957. (13) Brandes et al., 1959. (14) Electron microscopic studies of these viruses are going on in our laboratory. The hop virus is being studied by Nuber K (Landwirtschaftliche Hochschule, Hohenheim). (15) De Bruyn Ouboter et al., 1951. (16) Gold et al., 1957; Brandes, 1959. (17) Zimmer & Brandes, 1956. (18) Bode & Paul, 1956; Paul & Bode, 1956; Brandes & Paul, 1957. (19) Brandes, 1959. (20) These four legume viruses all have the same size (Brandes & Quanta 1955, Quanta 1959). Serological relationship has been established between bean common mosaic virus and bean yellow mosaic virus (Beenister & van der Want, 1951; Bereks, pers comm) and between bean yellow mosaic virus and pea mosaic virus (Goodchild. 1956). (21) Bode & Brandes, 1958. (22) Couch & Gold, 1954: 748 × 22 mm. Our determination of the NL revealed the same value (747 mm), but measurements in comnparison with TMV and other viruses revealed a diameter of about 12-13 μm (Brandes 1959). (23) This virus has been described and suspected of being related to sugar cane mosaic virus by Grancini (1957). It has been bee measured in our laboratory (Brandes 1959). (24) Gold et al., 1957. (25) Brandes & Zimnmer, 1955; Burghardt & Brandes, 1957.

Gemniniviruses (ジェミニウイルス)

Table 11. Geminiviruses(ssDNA) and their properties, distribution, and transmission. S: sedimentation coefficient. SCPS: Size of coat protein subunit

Name	Vector	Geographical distribution	In situ distribution	S	Nucleic acid		SCPS
					Type	Size
Maize streak	Cicadulina rnbila	Africa, India	-	76	ssDNA	7.1 × 105	28000
Bean golden mosaic	Bemisia lubaci	Tropical Am.	Phloem-associated parenchyma & sieve elements	69	ssDNA	8.0 × 105	27400
Cassava latent	-	Africa	-	76	ssDNA	8.0 × 105	34000
Chloris striate mosaic	Nesoclutha pallida	Australia	General but not epidermis	-	ssDNA	7.1 × l05	28000
Curly top	Circulifer tenellus	North Am.	Phloem-associated parenchyma	-	-
Euphorbia mosaic	Bemisia tabaci	Tropical Am.	Phloem-limited	-	-
Tomato golden mosaic	Bemisia tabaci	Tropical Am.	Phloem-limited	-	-
Tomato yellow dwarf	Bemisia tabaci	Japan	Phloem-limited	-	-
Tobacco leaf curl	Bernisia tabaci	Pantropical	-	-	-
Tobacco yellow dwarf	Orosius argentatus	Australia	-	76	-		27500
Bean summer death	Orosuis argentatus	Australia	Phloem-limited	-	-		-
Mungbean yellow mosaic	Bemisia tabaci	South Asia		-	-		-

Reovirus

Table 3. dsDNA segments and polypeptides of capsid proteins in plant reoviruses. #: number of RNA segment and polypeptide. PP: polypeptide. R: RNA segment (PTE) molecular weight (× 10⁶). MW_cal = calculated molecular weight (× 106). MW_obs = observed molecular weight (× 10⁶). No: Number of capsid proteins. Loc.: location (C = core, OL = outer layer, Ca = capsomere, OC = outer capsid, B = B-spike)

#	WTV					RDV					FDV	RBSDV	MRDV					PSV
		PP					PP							PP
	R	MWcal	MWobs	No	Loc	R	MWcal	MWobs	No	Loc	R	R	R	MWcal	MWobs	No	Loc	R
1	2.90	170.6	160.0	I	C	3.10	194.0	193.0	I	C	2.90	2.91	2.88	160.0
2	2.40	141.2	131.0	II	OL	2.50	156.0	152.0	II	OL	2.50	2.50	2.50	138.0	139.0	I	C	2.54
3	2.20	129.4	118.0	III	C	2.20	137.5	131.0	III	C	2.48	2.35	2.35	130.0	126.0	II	C	2.50
4	1.80	105.9				1.80	112.5				2.48	2.35	2.35	130.0	123.0	III	B	2.50
5	1.78	104.7	96.0	IV	OL	1.76	110.0	110.0	IV	OL	2.12	2.12	2.12	117.0	111.0	IV	OC	2.12
6	1.10	64.7				1.05	65.6				1.85	1.75	1.75	97.0	97.0	V	OC	1.75
7	1.05	61.8	58.0	V	C	1.02	63.7	62.0	V	C	1.45	1.45	1.45	80.0				1.48
8	0.83	48.8	36.0	VI	Ca	0.78	48.8	46.0	VI	OC	1.21	1.25	1.25	68.0				(1.25)
9	0.57	33.5	35.0	VII	Ca	0.70	43.8	45.0	VII	OC	1.15	1.18	1.18	65.0	64.0	VI	OC
10	0.55	32.4				0.67	41.9				1.12	1.10	1.08	60.0
11	0.54	31.7				0.48	30.0
12	0.32	18.8				0.48	30.0
Total	16.0					16.6					19.3	19.0	18.9					(19.6)

Transmission (伝搬)

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1. Contact (接触) 2. Seed transmission (種子) 3. Soil transmission (土壌) a. Nematode (線虫) Transmission of nepovirus and tobravirus through nematode Group cryptogram: [R/1:2.4/42/(1.4 – 2.2)/(27 – 40) – (2 × 1.4)/46:S/S:S/C, Ve/Ne Type member: tobacco ring spot virus = Main characteristics: Three types of isometric particle ca 28 nm in diameter with angular outlines, sedimenting at ca 50, 90-120 and 120-130 S and containing respectively ca 0, 27-40 and 42-46% single-stranded RNA. Two RNA species, M.W. ca 2.4 × 106 and 1.4-2.2 × 106, both necessary for infection. Each particle contains 60 molecules of a single coat polypeptide. M. Wt ca 55,000. Thermal inactivation point 55-70°C: longevity in sap a few days or weeks; concentration in sap 10-50 mg/l. Wide host range, causing ring spot and mottle symptoms, often with subsequent symptomless infection. Virus particles occur in cytoplasm, some in membranous tubules. Many cells contain a vesiculated cytoplasmic inclusion body. Transmissible by inoculation of sap, by soil-inhabiting nematodes and to progeny through seed and pollen. Table 8. Distribution of determinants for biological properties between the parts of virus genomes. RNA-1 a. RASPBERRY RINGSPOT VIRUS Seed transmissibility in Stellaria media Infectivity for raspberry cv. Lloyd George Suppression of symptoms in Petunia hybrida Lesion type in Chenopodium quinoa Systemic infection of Phaseolus vulgaris Systemic symptom severity in C. quinoa Speed of systemic symptom production in C. quinoa Competitiveness between virus genotypes s b. TOMATO BLACK RING VIRUS Seed transmissibility in S. media Speed of lesion production in C. quinoa Speed of systemic symptom production in C. quinoa c. TOBACCO RATTLE VIRUS ? RNA polymerase production Lesion type Lesion formation in P. hybrida Systemic infection of Nicotiana spp. RNA-2 a. RASPBERRY RINGSPOT VIRUS Coat protein specificity Vector specificity Yellowing symptoms in P. hybrida Lesion type in C. quinoa b. TOMATO BLACK RING VIRUS Coat protein specificity Vector specificity Lesion type in C. quinoa

c. TOBACCO RATTLE VIRUS Coat protein specificity Yellowing symptoms in Nicotiana spp. Length of short nucleoprotein particles b. Fungi (菌類) 4. Sap transmission (汁液) 5. Insect transmission (虫媒) Aphid (アブラムシ) Fig. Illustration of aphids in probing position. (A) Aphids on position to probe virus-infected tissues of the epidermis. (B) Aphids in feeding position, acquiring virus from phloem cells. Leaf beetle (ハムシ) = chrysomelid Tick (ダニ) Whitefly (コナジラミ) Issues 1. genome masking Table 6. General pattern of transmission (+) or non-transmission (-) of 5 isolates of barley yellow dwarf virus by 4 aphid species. Virus isolates R. padi M. avenae R. maidis S. graminum PAV + + - + RPV + - - + MAV - + - - RMV - - + - SGV - - - + → a mechanism that the early-infected virus protects against the infection of phylogenetically-different virus ↔ this virus does not have this mechanims = simultaneous replication Fig. 6. The combinations of proteins and nucleic acids extracted from the strains concomitantly infected by two virus species. C, D: transcapsidation, E, F: phenotypic mixing. 2. helper component

Viroids (ウイロイド)

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The smallest known pathogens, are naked, circular, single-stranded RNA molecules (200-400 bases) that do not encode protein yet replicate autonomously when infected into host vascular plants 1971 Diener, Theodor Otto (1921-2023, Swiss-American) discovered potato spindle tuber viroid (PSTVd, ジャガイモやせ芋ウイロイド) = 130000 Da → single-stranded circular RNA (sscRNA) ☛ RNA world 1979 ASBVd: characteristics different from PSTVd Taxonomy Nomenclature: described in English (not Latin) host plant + characterized desease + viroid Similarity of genome sequence > 90% ⇒ same species varieteis when needed, even if siilarity > 90% ≥ 30 viroids Family Avsunviroidae Avsunviroid: Avocado sunblotch viroid (ASBVd) Pelamoviroid: Chrysanthemum chlorotic mottle viroid (キク退緑斑紋ウイロイド), Peach latent mosaic viroid (モモ潜在モザイクウイロイド) Elaviroid: Eggplant latent viroid

Family Pospiviroidae Pospiviroid: Chrysanthemum stunt viroid (キク矮化ウイロイド), Citrus exocortis viroid (カンキツエクソコーティスウイロイド), Columnea latent viroid, Iresine viroid 1, Mexican papita viorid, Pepper chat fruit viroid, Potato spindle tuber viroid, Tomato apical stunt viorid, Tomato chlorotic dwarf viroid (トマト退緑萎縮ウイロイド), Tomato planta macho viroid Hostuviroid: Hop stunt viroid (ホップ矮化ウイロイド) Cocadviroid: Citrus bark cracking viroid (カンキツバーククラッキングウイロイド), Coconut cadang-cadang viroid, Coconut tinangaja viroid, Hop latent viroid (ホップ潜在ウイロイド) Apscaviroid: Apple dimple fruit viroid (リンゴくぼみ果ウイロイド), Apple scar skin viroid (リンゴさび果ウイロイド), Australian grapevine viroid (ブドウオーストラリアウイロイド), Citrus bent leaf viroid (カンキツベントリーフウイロイド), Citrus dwarfing viroid (カンキツ矮化ウイロイド), Citrus viroid V (カンキツウイロイドV), Citrus viroid VI (カンキツウイロイドVI), Grapevine yellow speckle viroid 1 (ブドウ黄色斑点ウイロイド1), Grapevine yellow speckle viroid 2, Pear blister canker viroid (ナシブリスタキャンカーウイロイド) Coleviroid: Coleus blumei viroid 1 (コリウスウイロイド1), Coleus blumei viroid 2, Coleus blumei viroid 3 Disease symptom of host Stem and leaf: mottle, stunt, dwarf and yellow spot Woody plants: mottle, epinasty (葉下垂), exocortis (樹皮剥離)