• TABLE OF CONTENTS
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 Front Cover
 Title Page
 Table of Contents
 List of Tables
 Introduction
 Virus genome control of cylindrical...
 Cylindrical inclusions in classification...
 Problems and advantages involved...
 Some comments on assignments of...
 Summary and conclusions
 Reference
 Figures
 Tables
 Back Cover














Group Title: Bulletin - University of Florida. Agricultural Experiment Station ; no. 894
Title: Cylindrical inclusions
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 Material Information
Title: Cylindrical inclusions
Series Title: Bulletin
Physical Description: 79 p. : ill. ; 28 cm.
Language: English
Creator: Edwardson, J. R ( John Richard ), 1923-
Christie, R. G
Publisher: University of Florida, Agricultural Experiment Station, Institute of Food and Agricultural Sciences
Place of Publication: Gainesville FL
Publication Date: [1996]
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Subject: Potyviruses   ( lcsh )
Potyvirus diseases   ( lcsh )
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Bibliography: Includes bibliographical references (p. 13-41).
Statement of Responsibility: J.R. Edwardson and R.G. Christie.
General Note: "May 1996".
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Table of Contents
    Front Cover
        Page i
    Title Page
        Page ii
    Table of Contents
        Page iii
    List of Tables
        Page iv
    Introduction
        Page 1
        Page 2
    Virus genome control of cylindrical inclusions
        Page 3
        Page 4
    Cylindrical inclusions in classification and diagnosis
        Page 5
        Page 6
    Problems and advantages involved in using subdivisions in diagnosis
        Page 7
        Page 8
    Some comments on assignments of viruses to subdivisions
        Page 9
        Page 10
    Summary and conclusions
        Page 11
        Page 12
    Reference
        Page 13
        Page 14
        Page 15
        Page 16
        Page 17
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    Figures
        Page 43
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        Page 46
        Page 47
        Page 48
        Page 49
        Page 50
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        Page 52
    Tables
        Page 53
        Page 54
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    Back Cover
        Back cover
Full Text
100
y 1o

May 1996


Cylindrical Inclusions





J. R. Edwardson

R. G. Christie


Marston Science
Library
OCT 101996
University of Florida


UNIVERSITY OF
FLORIDA


Agricultural Experiment Station
Institute of Food and Agricultural Sciences


Bulletin 894




















Cylindrical Inclusions

J.R. Edwardson and R.G. Christie





















J.R. Edwardson is professor and R.G. Christie is senior biologist, Department of Agronomy, Institute of Food and
Agricultural Sciences, University of Florida.










CONTENTS
C hap ter 1. In trodu action ....................................... ........................................................ ...............................1
Chapter 2. Virus Genome Control Of Cylindrical Inclusions ..................................................3
Chapter 3. Cylindrical Inclusions In Classification And Diagnosis ............................................. ....... 5
Chapter 4. Problems and Advantages In Using Subdivisions In Diagnosis ..............................................7
Chapter 5. Some Comments On Assignments Of Viruses To Subdivisions .............................................9
Chapter 6. Sum m ary And Conclusions ...................................................... .... ..... ........................... 11
R eferen ces ................... . .......................................................................................................................................13



FIGURES
Figure 1. Pinwheels induced by Nothoscordum Mosaic and Potato Virus-Y Potyviruses ..................43
Figure 2. Pinwheels and bundles induced by Wheat Streak Mosaic Rymovirus ..............................43
Figure 3. Cylindrical inclusion components induced by Henbane Mosaic Potyvirus ......................44
Figure 4. Type-1 cylindrical inclusions induced by Papaya Ring-spot-W Potyvirus .........................44
Figure 5. Scrolls induced by Blackeye Cowpea Mosaic Potyvirus .................................... ..... ...44
Figure 6. Type-2 cylindrical inclusions induced by Lettuce Mosaic and Bean Yellow Mosaic
P oty v iru ses .............. ............................................................. ...................................... ................45
Figure 7. Type-2 cylindrical inclusions induced by Tobacco Etch Potyvirus ...................................45
Figure 8. Type-4 cylindrical inclusions induced by Potato Virus-Y and Turnip Mosaic
Potyviruses ............................ ....... .... ......... ... ... ... ..... ... ................... ....46
Figure 9. Type-2 cylindrical inclusions induced by Tobacco Etch Potyvirus............................... ....46
Figure 10. Laminated aggregates induced by Tobacco Etch and Bidens Mottle Potyviruses ..............47
Figure 11. Laminated aggregates induced by Tobacco Etch Potyvirus........................... .... ...47
Figure 12. Portion of a cylindrical inclusion exhibiting striations induced by Potato Virus-Y
P otyv iru s ........................................ ........ ............ ... .. .. ... ... ........ ... ................48
Figure 13. Striations in cylindrical inclusions induced by Nothoscordum Mosaic Potyvirus and
W heat Streak M osaic Rym virus .................................................................................. 49
Figure 14. Striations in cylindrical inclusions induced by Desmodium Mosaic Potyvirus ...................49
Figure 15. Type-1 cylindrical inclusions induced by Papaya Ring- spot-W Potyvirus and type-3
cylindrical inclusions induced by Turnip Mosaic Potyvirus ............................. .............50
Figure 16. Cylindrical inclusions induced by an uncharacterized Potyvirus in garlic tissues ..............51
Figure 17. Cylindrical inclusions induced by Moroccan Watermelon Mosaic Potyvirus ......................52
Figure 18. Type-3 and type-4 cylindrical inclusions induced by Watermelon Mosaic Virus-II
P otyv iru s ................................................................................................ ................................... 52










TABLES


Table 1. Cytoplasmic Cylindrical Inclusions (CI) Induced by Viruses of the Family Potyviridae,
Genus Potyvirus (Aphid-Transmitted) ................................................................................................53


Table 2. Cytoplasmic Cylindrical Inclusions Induced by Viruses of the Family Potyviridae, Genus
Rymovirus (Eriophyid Mite-Transmitted) ..............................................................................................68


Table 3. Cytoplasmic Cylindrical Inclusions Induced by Viruses of the Family Potyviridae, Genus
Bym virus (Fungus- Transm itted) ......................................................................................................... 69


Table 4. Viruses Exhibiting Potyvirus Properties but Which Are Presently Unassigned to the Family
Potyviridae ....................................................................................................................................... .................70


Table 5. Viruses Whose Cylindrical Inclusions Have Not Been Characterized ...................................... ..71


Table 6. Viruses Inducing Type-1 Cylindrical Inclusions Constitute Subdivision I ...........................................72


Table 7. Viruses Inducing Type-2 Cylindrical Inclusions Constitute Subdivision II....................................... 74


Table 8. Viruses Inducing Type-3 Cylindrical Inclusions Constitute Subdivision III .........................................76


Table 9. Viruses Inducing Type-4 Cylindrical Inclusions Constitute Subdivision IV ................................... 77


Table 10. Viruses of the Potyviridae with no Cytological Studies ......................................................................79








CHAPTER I

Introduction

Cylindrical inclusions (CIs) are proteinaceous,
striated sheets in various configurations, induced in the
cytoplasm of host plants by viruses of the Potyvirus,
Rymovirus and Bymovirus genera of the family
Potyviridae. While plant viruses in all the other 34 recog-
nized groups also induce inclusions, none induces cylin-
drical inclusions (Edwardson et al. 1993). CIs have been
observed in all plant cell types but they occur frequently
in epidermal and mesophyll cells.
Although groups of CIs were detected by light
microscopy in the 1920s (Holmes, 1928; Kunkel, 1922),
the shapes of individual CIs were not understood until the
1960s (Edwardson, 1966a,b; Rubio-Huertos and Lopez-
Abella, 1966). Electron microscopy of thin-sections
reveals that the central portions of CIs consist of thin,
more or less curved plates (arms) with sides converging
around a central core and opposite sides diverging to form
pinwheels in cross sections (Figure la,b). In longitudinal
sections the thin plates appear as groups of parallel lines
which are described as bundles (Figure 2). In some
sections the cylindrical inclusion plates may not be
oriented in pinwheel-bundle configurations, but appear as
a tangled mass (Figure 3). In some virus-infections the
pinwheel arms may extend and roll up into scrolls which
appear as circles or coils in cross sections (Figure 4). In
longitudinal sections the scrolls appear as tubes (Figures
5,17). In other virus infections the pinwheel arms may
extend and in cross sections appear as straight flat plates
(Figures 6a, 7) or more or less curved plates (Figure 6b).
These plates are termed laminated aggregates because of
their layered appearance. These structures can be recog-
nized in cross sections when attached to the central
portion of the CI (Figure 9) and in longitudinal sections
where the CIs central portions do not appear (Figure
10a,b). The layered structure of laminated aggregates is
not apparent in tangential sections such as those in the
upper right portion of Figure 11.


When CIs are examined in negative stain in the
electron microscope, all of their components exhibit
striations with a periodicity of ca 5 nm (Figure 12). The
striations are also apparent in thin sections of vapor-fixed,
vacuum-dehydrated tissues (Edwardson et al., 1968).
These treatments produce a reversal of contrast to images
arising from conventionally prepared sectioned tissues
(Figures 4, 7).
The vapor-fixation, vacuum-dehydration tech-
nique has been useful in detecting substructure of CIs in a
number of plant species (Figures 4, 7, 13). Reversed
contrast of images of CIs has been observed occasionally
in conventionally prepared thin sections, such as in
Desmodium canum infected with Desmodium mosaic
Potyvirus (Figure 14).
At one time many plant virologists regarded
cylindrical inclusions and other types of inclusions as
either by-products of altered metabolism of infected cells
or as accumulations of virus particles. It has been demon-
strated that many viruses in many groups do induce
inclusions consisting of virus particles (Christie and
Edwardson, 1977; Martelli and Russo, 1977). However,
cytological and serological studies have demonstrated that
CIs are not composed of virus particles (Figures 7, 12, 13,
14), and biochemical investigations indicate that CIs
possess RNA-helicase activity (Lain et al. 1989, 1990).
Serological relationships have been established between
CIs induced by many members of the Potyvirus genus.
Also a serological relationship between CIs induced by
tobacco etch Potyvirus and wheat streak mosaic
Rymovirus has been demonstrated (Purcifull and Hiebert,
1992).
This report describes the different types of
cylindrical inclusions and explains how one type might be
interpreted as being another type when only limited
cytological information is available. Members of the
Potyviridae have been separated into Subdivisions on the
basis of the type of CI each virus induces. Determining
the type of CI should allow the investigator to assign the
virus to a Subdivision and thereby reduce the number of
comparisons necessary for identifying the virus.











CHAPTER II
Virus Genome Control of
Cylindrical Inclusions

That CIs are controlled by portions of the virus
genome has been demonstrated in the following ways:

1) A virus in the family Potyviridae induces the
same type of CI in different hosts (these can include
species, genera, families and classes) (Edwardson, 1974);

2) Different viruses induce their characteristic
types of CIs in the same host (Edwardson, 1974);

3) CI protein is serologically unrelated to virus
and host proteins (Purcifull et al. 1973);

4) Potyvirus RNA has been translated in vitro in
rabbit reticulocyte systems; one of the translation products
is CI protein (Dougherty and Hiebert, 1980);

5) Viruses in other groups induce inclusions, but
not CIs (Barnett, 1991; Edwardson et al. 1993).


Inclusions were considered as one of dozens of
criteria for use ih classifying plant viruses by the Interna-
tional Committee On Nomenclature of Viruses (Harrison
et al. 1971; Wildy, 1971). But these taxonomists did not
use information about inclusions in classifying any virus
in their 16 groups. However, a few years later, in 1976,
CIs were included as a main characteristic of the
potyvirus group by the International Committee On
Taxonomy Of Viruses (ICTV) (Fenner, 1976), and they
remain a main characteristic of the family Potyviridae
(Barnett, 1991). The viruses in each genus of the family
induce CIs. Harrison and coworkers (1971) defined a
plant virus group as "a collection of viruses and/or virus
strains each of which shares with the type member all or
nearly all of the main characteristics of the group. These
distinguishing main characteristics can be found only by
considering the information available on as many as
possible of the properties of each virus. We feel that
failure of a virus to share one or a few main characteris-
tics with the type member of a virus group should not
exclude it from membership of that group. However, most
properties should be shared".











CHAPTER III

Cylindrical Inclusions in

Classification and Diagnosis

At present the ICTV lists 182 viruses in the family
Potyviridae including 166 viruses in the genus Potyvirus,
7 in the Rymovirus and 5 in the Bymovirus genera
(Barnett, 1991; Pringle, 1993). All viruses in the family
which have been studied cytologically induce CIs. Two
viruses included in the Family Potyviridae (Maclura
mosaic and Narcissus latent) are listed as aphid transmit-
ted but are unassigned to a genus, and two viruses (sweet
potato mild mottle and sweet potato yellow dwarf) listed
as whitefly-transmitted are also unassigned to a genus in
the Potyviridae (Barnett, 1991). These unassigned viruses
induce CIs.
We could not find reports of cytological studies on
31 of the species and tentative species in the Potyvirus genus
(Tables 1,10). One hundred and thirty-five species and
tentative species of the genus Potyvirus have been studied
cytologically, and all induce CIs (Table 1). The 5 species and
2 tentative species in the Rymovirus genus (Table 2) and the
5 species in the Bymovirus genus also induce CIs (Table 3).
The viruses in the 3 genera which constitute the family
Potyviridae have many characteristics in common: particle
morphology, physicochemical properties, nucleic acid,
protein and induction of CIs (Barnett, 1991, 1992;
Edwardson, 1992a). Some species of the Rymovirus
(Lesemann and Vetten, 1985; Shukla et al. 1989) and
Bymovirus (Stanarius et al. 1989) genera are serologically
related to some Potyvirus species. The viruses comprising
the genus Potyvirus are aphid transmitted, those of the genus
Rymovirus are mite transmitted, and viruses of the genus
Bymovirus are fungus transmitted. Bymoviruses are bipartite
while viruses of the other genera possess monopartite
genomes.
Seventeen viruses in the Potyviridae induce CIs
which have not yet been characterized (Table 5). Each of the
viruses listed in Table 5 has been examined cytologically;
most of the reports involve electron microscopy, and they
describe parts of the central portions of CIs (pinwheels and
bundles) such as those in Figures 1 and 2. The information
in these reports is insufficient for assigning viruses to subdi-
visions.
Fifteen viruses possessing some of the characteris-
tics of potyviruses but which are as yet unassigned to the
Family Potyviridae by the ICTV are listed in Table 4.
Fourteen of these viruses have been studied cytologically
and each induces CIs. For those instances in which cyto-
logical information is adequate, viruses of the Potyviridae


have been separated into 4 Subdivisions (Edwardson,
1974; Edwardson and Christie, 1991a; Edwardson et
al.1984). Each of the Subdivisions contains viruses and
strains inducing a particular type of cylindrical inclusion.
Subdivision-I is made up of 49 viruses which induce only
type-1 CIs (Table 6). Type-1 CIs consist of the central
portion (pinwheels and bundles) with scrolls (circles or
coils in cross section) as in Figure 15a and tubes (in
longitudinal section) as in Figure 5. The scrolls may
appear attached to the central portion of the CI (as in
Figure 4), or unattached (as in Figures 5 and 17). Many
viruses in the Potyvirus and Rymovirus genera induce
type-1 CIs. Of the 49 viruses in Subdivision-I, 30 are listed
as being tentatively assigned to the Subdivision, and they
are designated by (I). These latter assignments are based
on only single reports.
Viruses assigned to Subdivision-II (Table 7)
induce only type-2 CIs consisting of pinwheels, bundles
and more or less straight long laminated aggregates
(Figures 6a, 7, 9, 10, 11), and pinwheels, bundles and more
or less curved long laminated aggregates (Figure 6b).
Twenty-six viruses have been assigned to this Subdivision
as members and 19 as tentative members. These latter
assignments which are designated by (II) are based on only
single reports.
The 19 viruses and 8 virus strains assigned to
Subdivision-III induce type-3 CIs which consist of pin-
wheels, bundles, scrolls and more or less straight laminated
aggregates (Figure 15b). Some viruses in the Potyvirus,
Bymovirus and Rymovirus genera induce type-3 CIs (Table 8).
The 35 viruses and 7 virus strains listed in Table 9
induce type-4 CIs which consist of pinwheels, bundles,
scrolls and short, usually curved, laminated aggregates
(Figure 8a,b). In our experience the scrolls usually outnumber
the laminated aggregates.
Both scrolls and laminated aggregates may attain
considerable lengths so that they are often observed in no
apparent association with the central portion of the CIs.
These conditions are illustrated in Figures 10 and 17.
Most investigations of CIs have used electron
microscopy of thin sections. Unless electron microscopy is
carried out in conjunction with light microscopy, much
time and effort can be devoted to studying thin-sections
which do not contain CIs. The electron microscope has a
very limited sampling capacity when thin-sections rather
than extracts are investigated. Small samples of tissue
(usually ca 1 X 3 mm) are dehydrated, fixed, stained,
embedded, mounted, trimmed and sectioned. Useable
sections are ca 600 nm or less in thickness. Only limited
areas of these small samples are available for study since
parts of the sections are blocked out by the supporting grid
mesh. At the time of sampling some CIs may not have







developed components beyond those comprising the
central portion (pinwheel in cross section, bundle in
longitudinal section). When all CI components have
developed, limited sampling can result in misunderstand-
ing of the inclusion's morphology which can lead to
misassignments of viruses to Subdivisions.
Limited sampling of thin-sections of infected
tissues may result in failure to detect the presence of CIs or
to misidentification of CI types. Viruses and the inclusions
they induce are not uniformly distributed in host tissues.
Inclusions are not static; they increase in number and size
within days to weeks after infection. In early infections,
small CIs develop at the plasmalemma and are so small
that their detection is often difficult. Sampling tissues from
portions of the host with long-standing infections may
result in examining the debris of inclusions which are
degenerating. CIs are three-dimensional structures, and
different aspects of these structures will appear in different
sectional planes. All the components of CIs may not
appear in the same section. Therefore, examination of a
limited number of sections may provide only partial
information on the structure of the CIs. For example, some
wheat streak mosaic Rymovirus isolates induce type-3 CI
(Gardner, 1973) while others induce type-4 (Baker et al.
1985). However, some sections may exhibit only pin-
wheels and bundles (Figure 2a) while some sections show
the presence of pinwheels and scrolls (Figure 13b). There-
fore, Figures 2a and 13b contain incomplete representa-
tions of type-3 or type-4 CIs. Another example of prob-
lems arising from examining only a few sections is pre-
sented in Figure 16a and b. Here sections of Allium
sativum infected with a potyvirus were obtained from
different parts of the same leaf. If only Figure 16a were
considered, the type-1 CIs there would suggest assignment
of the virus to Subdivision-I. However, the CIs in Figure
16b contain scrolls and laminated aggregates.
Some of the viruses presently assigned to Subdivi-
sion-I (Tables 1, 6) may actually induce type-3 or type-4
CIs, but because sampling has been inadequate to reveal
all the CI components, the viruses appear to induce only
type-1 CIs. Also, some of the viruses presently assigned to
Subdivision-II (Tables 1, 7) may induce type-3 or type-4
CIs, but because sampling has been inadequate to reveal


all the components of the CIs, the viruses appear to induce
only type-2 CIs. Viruses which have been subjects of
several cytological studies seem less likely to be
misassigned to Subdivisions than those less studied. Many
reports cited in Tables 1-4 provide only limited informa-
tion on CI structure. Extensive cytological investigations
of inclusions were not the principal objective of most of
the reports in Table 1. Many of the authors of these reports
used the presence of CIs to establish that their viruses were
members of the Family Potyviridae.
The cylindrical inclusions are very interesting
complex structures. They are also of practical value to
taxonomists in that their presence indicates the presence of a
virus belonging to the largest of the plant virus families,
the Potyviridae. Cylindrical inclusions are useful to
diagnosticians in that the different morphologies of these
inclusions have led to construction of Subdivisions.
Determining that a virus is a member of a Subdivision
reduces significantly the number of comparisons required
to establish the virus as a species or strain.
The structure of cylindrical inclusions has been
established by electron microscope studies of inclusions in
extracts and in thin-sections. Differences in CI morpholo-
gies have been established through electron microscopy.
The different types of CI have also been studied in the
light microscope (Christie and Edwardson, 1977, 1986;
Edwardson and Christie, 1991; Edwardson et al. 1993).
The CIs are green in the combination protein-stain
Calcomine Orange-Luxol Brilliant Green, and are un-
stained in the nucleoprotein-stain Azure A (Christie and
Edwardson, 1977, 1986). Individual pinwheel arms are not
resolved in the light microscope; however, scrolls appear
as dots when they are viewed end-on and remain as dots
when the focus is changed. Scrolls appear as lines in side-
view and they disappear rapidly as focus is changed.
Laminated aggregates appear as lines and remain as lines
when the focus is changed. Cylindrical inclusion types 1,
2, and 3 can be readily distinguished in the light micro-
scope. The short, curved, laminated aggregates of the type-
4 cylindrical inclusions are often difficult to detect.
Therefore, type-1 and type-4 CIs are often difficult to
distinguish (Christie et al. 1995).








CHAPTER IV

Problems and Advantages

Involved in Using Subdivisions in

Diagnosis

Subdivisions were not constructed with any
anticipation of their use in virus taxonomy. Separations of
viruses on the basis of CI morphology do not coincide in
any discernable way with separations based on vectors,
serology or host ranges (Edwardson, 1992a). That the
presence of CIs is diagnostic for infections by viruses of
the family Potyviridae is well established. Subdividing the
Potyviridae on the basis of CI structure for purposes of
identification has proven value (Lesemann, 1988). Using
differences in CI morphology to form Subdivisions was
undertaken to assist in separating virus species. Reducing
the number of comparisons necessary for virus identifica-
tion such as in host ranges and serological tests assists the
diagnostic process.
Different strains and isolates of the same virus
induce a single type of CI, such as those of artichoke
latent, bean yellow mosaic and cowpea aphid-borne
mosaic viruses (Table 1). However, different strains and
isolates of the same virus may induce different types of
CI, such as those of sugarcane mosaic, maize dwarf
mosaic and peanut mottle viruses (Table 1). This latter
situation is expected. Portions of the genomes of viruses
in the family Potyviridae control cylindrical inclusion
formation, and mutations would be expected in these as
well as other portions of the genomes.
Prior to 1984 we assumed that potato virus Y (PVY)
induced only type-4 CIs and that reports of PVY inducing
only type-1 or only type-2, or only the central portions could
all be accounted for by assuming inadequate sampling.
However, extensive investigations of an Australian strain of
PVY (Moghal and Francki, 1981; Edwardson et al. 1984;
Francki et al. 1985) indicate that this Australian strain induces
only type-1 CIs.


When viruses which induce type-3 and others
which induce type-4 CIs are also reported to induce type-
1 or type-2 inclusions, it has been assumed that the type-1
and type-2 inclusions represent portions of type-3 or type-
4 CIs (Table 1). This assumption appears to be well
substantiated by the CIs reported for turnip mosaic
(McDonald and Hiebert, 1975; lino et al. 1987), garlic
mosaic (Chang et al. 1988), carnation vein mottle
(Paludan and Begtrup, 1974; Begtrup, 1976) and
passionfruit woodiness (Teakle and Pares, 1977), where
CIs of different types are shown to occur in the same host
tissues and in some cases in the same cell. However, some
of the reports of CIs induced by soybean mosaic strains
and isolates are not readily accounted for by this assump-
tion. The instances of type-1 and type-3 CIs (lizuka and
Yunoki, 1975) and type-1 and type-4 CIs (Nicolaescu et
al. 1976; Hunst and Tolin, 1983) occurring in the same
tissues appear to support the assumption. However, when
sampling is used to explain studies describing only type-1
inclusions where types-3 or -4 are expected, then sam-
pling should also result in some reports) of only type-2
CIs being induced by soybean mosaic virus. We have been
unable to find such an example. Therefore, we assume
that some strains of soybean mosaic virus induce type-1
CIs while others induce type-3 and others type-4 CIs.
Applications of cytological studies to diagnosis of
infections by members of the family Potyviridae at the
family level are not complicated by the fact that some
strains of a virus induce different types of CIs. But, the
occurrence of such strains makes more difficult the
application of information on Subdivisions to diagnosis at
the species level. Information on types of CIs is yet to be
obtained for many members of the Potyviridae (Tables 1,
5, 10). The usefulness of Subdivisions for assisting
diagnosis of virus infections at the species leyel will be
much improved as cytological information on these
viruses becomes available.











CHAPTER V

Some Comments on Assignments

of Viruses to Subdivisions

There may be exceptions to the statement in
Chapter IV that proclaimed only a single type of CI is
induced by individual strains and isolates of the same
virus. For example, the blackeye cowpea mosaic strain of
bean common mosaic virus (BCMV) has been reported to
induce only type-1 CIs, while the peanut stripe strain of
BCMV is reported to induce type-4 inclusions (Table 1).
However, sugarcane mosaic virus strain H (now included
in the sorghum mosaic virus species by Lesemann et al.
1992) has been shown to induce different types of CIs:
type-3 (Edwardson, 1974), type-4 (Lesemann et al. 1992)
and type-1 (Ford, 1968; Giorda et al. 1986; Hearon et al.
1981). While the type-1 CIs may be explained by assum-
ing they arise from samplings from type-3 and/or type-4
inclusions, the existence of both type-3 and type-4 CIs in
the H strain can be interpreted in different ways:

1) The gene controlling CI induction in the H
strain has mutated resulting in some H strain isolates
inducing type-3 CIs, other H strain isolates inducing type-
4 CIs.;

2) The H strains examined by Lesemann et al.
(1992) and Edwardson (1974) are different strains or
different viruses;

3) The gene controlling CI formation in strain H
induces more than one type of CI.

The majority of cytological studies of celery
mosaic virus (CeMV) infections have reported type-1 CIs
(Table 1). Buturac et al. (1974) examined a CeMV isolate
which induced type-1 CI and did not infect Ammi majus.
Their isolate is serologically related to but is not identical
to Italian, English and East German "celery mosaic virus."
The isolates Ag, Dc, Pc and P described by Horvath et al.
(1976) are serologically identical to Italian "celery mosaic
virus." The P isolate does infect Ammi majus and it
induced type-2 CIs. It appears that there are two
potyviruses in Europe which are different cytologically
and serologically; however, both are named celery mosaic
virus.
Malva vein clearing virus (MVCV) has been
described as inducing type-1 CIs (Table 1). Horvath et al.
(1979) reported the induction of type-3 cylindrical inclu-
sions in Malva silvestris by Hungarian and Yugoslavian


isolates of a virus isolated from naturally infected Malva
sp. plants. These isolates infected only members of the
Malvaceae, however, MVCV infects species in the
Malvaceae, Leguminosae and Solanaceae. No serological
comparisons have been conducted between MVCV and the
virus isolates described by Horvath et al. (1979). Henriques
and Henriques (1986) also reported the occurrence of type-3
cylindrical inclusions in field-collected leaves of a Malva sp.
which exhibited vein clearing symptoms. Whether this
Canadian potyvirus and those studied by Horvath et al.
(1979) are MVCV remains to be determined.
The occurrence of type-1 CIs (McWhorter, 1937;
Edwardson, 1974) and type-2 CIs (Takahashi, 1973) in onion
yellow dwarf virus (OYDV) infected tissues (Table 1) might
be accounted for by assuming that:

1) Some strains of OYDV induce type-1, others
type-2 CIs;

2) Different potyviruses are named "onion yellow
dwarf', one induces type-1 and the other induces type-2 CIs;

3) OYDV induces type-3 or type-4 CIs, and the
reported type-1 and type-2 inclusions are the result of
inadequate sampling.

Cytological investigations of tissues infected with
papaya ringspot virus (P-isolates) have, with one exception,
described only type-1 CIs. The exception is the report of
Chen (1984) in which type-1 inclusions are described and in
addition some pinwheel arms are interpreted as laminated
aggregates (LA). We assume that the LAs in Chen's (1984)
report are actually pinwheel arms which have been misinter-
preted as LAs, resulting in the misassignment of Chen's
isolate to Subdivision-III.
Martelli and Russo (1976) reported that a virus
inducing type-1 CIs in Cucurbita pepo was watermelon
mosaic virus-2 (WMV-2). Some strains of WMV-2 induce
type-3, others induce type-4 CIs (Table 1). However,
Purcifull and Hiebert (1979) demonstrated that Martelli and
Russo's virus was serologically identical to papaya ringspot-
W isolates (formerly watermelon mosaic virus-1). Papaya
ringspot-W induces type-1 CIs (Table 1).
Most peanut mottle virus isolates have been shown
to induce type-2 CIs while two isolates are reported to
induce type-3 CIs (Table 1). Gourett-Triharso (1977)
reported type-4 CIs in peanut tissues infected with "Arachis
Mottle" virus. Other than being a potyvirus, the relationship
of this virus to peanut mottle virus is unknown.
Different viruses seem to have been described
under the name pea seed-borne mosaic virus (PSBMV).
At present it is not possible to assign the virus to a Subdi-







vision. Hampton et al. (1973) have reported that in Pisum
sativum PSBMV induced types-1, -2, -3 and -4 CIs and
crystalline cytoplasmic inclusions which are unlike those
induced by bean yellow mosaic virus. Tissues infected
with the S, isolate contained CIs but also virus aggregates
and vesiculated cytoplasm which were characteristic of
Carlavirus infections. Pisum sativum infected with the C4
isolate of PSBMV exhibited cytoplasmic crystalline
inclusions of the kind induced by bean yellow mosaic
virus. Before ending this chapter on the assignment of
viruses to Subdivisions, the question of the number of
Subdivisions should be addressed. We know from our
own studies and from the investigations of many others
that there are four types of cylindrical inclusions and that
the viruses inducing a certain type of CI can be placed in a
certain Subdivision. Are there more than 4 types of CI and
more than 4 Subdivisions? We do not know, but the
answer may be yes.
There may be viruses which induce CIs consisting
only of the central portion, plates which form pinwheels
in cross section, bundles in longitudinal section without


any scrolls or laminated aggregates. The CIs could look
something like those in Figures 1 and 2a. However, the
CIs in Figures la,b and 2a do not represent a new type;
they were induced by Nothoscordum mosaic and potato
virus Y Potyviruses and wheat streak mosaic Rymovirus,
respectively. Viruses inducing the "new" type CIs would
form another Subdivision.
There may be viruses capable of inducing both
type-3 and type-4 CIs (not induced by different strains of
the same virus but by a single strain). It has been assumed
that different strains of the same virus have induced types
3 and 4 CIs in tissues infected with wheat streak mosaic
Rymovirus (McMullen and Gardner, 1980); and by
Potyviruses leek yellow stripe (Horvat and Verhoyen,
1975), tobacco vein banding (Reddick et al. 1992) and
turnip mosaic (Stefanac and Wrischer, 1989). However, a
single strain of these viruses with a CI-controlling gene of
a different pleiotropic ability could account for the above
reports. Inclusions induced by such a virus may appear
similar to those in Figures 16 and 18. This and other CI
types may eventually be established as the bases of
additional Subdivisions.








CHAPTER VI

Summary and Conclusions


Potyviruses induce cylindrical inclusions (CIs) of
4 recognized types in host cytoplasm. Viruses inducing
certain types of CI have been combined in 4 subdivisions
of the family Potyviridae: Subdivision-I presently consists
of 49 viruses inducing type-1 inclusions (pinwheels,
bundles and scrolls); Subdivision-II contains 45 viruses
inducing type-2 inclusions (pinwheels, bundles and
laminated aggregates); Subdivision-III contains 27 viruses
inducing type-3 inclusions (pinwheels, bundles, scrolls
and laminated aggregates); 42 viruses inducing type-4 CIs
(pinwheels, bundles, scrolls and short usually curved
laminated aggregates) comprise Subdivision-IV. The
possibility of the existence of additional types of CIs is
discussed. The use of Subdivisions to separate this largest
of plant virus families, the Potyviridae, assists in the
diagnosis of infections induced by the viruses of the
family. Applying information on morphology of CIs to
diagnosing virus infections is carried out in many labs,
although the applications are complicated by different
strains of the same virus inducing different CI types, and
are hampered by lack of information about CIs induced by
many viruses in the Potyviridae. To the workers in such
labs we wish continued success in applying cytology to
virus disease diagnosis.


More cytological investigations of viruses pres-
ently assigned as tentative members of Subdivisions
(Tables 1-9), and of those viruses in the Potyviridae which
have not been studied cytologically (Tables 1, 10) or
whose CIs have been only partially described (Table 5)
will improve the usefulness of the Subdivisions in diagno-
sis of virus infections at the species level.
Cytological investigations of virus infections have
also indirect applications to diagnosis of specific diseases.
No culture collection or laboratory possesses antisera to
the hundreds of plant viruses comprising the 35 groups
now recognized by the ICTV. Inclusions are diagnostic for
infections induced by most plant viruses at the group level
(Christie and Edwardson, 1977; Edwardson et al. 1993).
Determining the group to which a virus belongs reduces
the number of serological tests which will be necessary to
identify the virus. When a diagnostician's cytological
investigations reveal CIs, infection by a member of the
Potyviridae is established. Ascertaining the Subdivision
assignment of a Potyviridae virus reduces the number of
tests required for virus identification at the species level.


Summary


Conclusions











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mosaic virus infecting pea (Pisum sativum). Jiangsu
Journal Agricultural Science 7:11-17.








,-,
k ... "


Figure lb. Pinwheels induced by
Potato Virus Y Potyvirus in Nicotiana
tabacum NN. X 27,700



.
A ;* -.''. *'*


Figure 2. Pinwheels (center) and
bundles (longitudinal sections of
cylindrical inclusions) (upper left)
induced by Wheat Streak Mosaic
Rymovirus in Triticum aestivum. X
47,800


At' '' '


'.y ; ,r,.'
",i,'r

Jl
` kr


tL.4


Figure la. Pinwheels (cross-sections
of cylindrical inclusions) induced by
Nothoscordum Mosaic Potyvirus in
Nothoscordumfragrans. Black dots are
cross sectioned virus particles. Sev-
eral pinwheels have virus particles at
their centers. X 47,800













Figure 3. Cylindrical inclusion arms,
most of which are not organized into
pinwheels and bundles, induced by
Henbane Mosaic Potyvirus in Nicoti-
ana tabacum X 27,700








Figure 4. Papaya Ringspot-W
Potyvirus (formerly watermelon
mosaic virus-1) induced type-1
cylindrical inclusion (pinwheel and
scroll) in Cucurbita pepo. Vacuum
dehydration, OsO4 vapor fixation.
Note striations on cylindrical inclu-
sion components, and note in the
center of the figure four cross-sec-
tioned virus particles (arrow).
X 180,000









Figure 5. Tubes (scrolls in longitudi-
nal-section) induced by Blackeye
Cowpea Mosaic Potyvirus in Vigna
unguiculata. X 47,800







Figure 6a. Type-2 cylindrical inclu-
sions (pinwheels with straight arms
and laminated aggregates) induced
by Lettuce Mosaic Potyvirus in
...*' Lactuca sativa. X 27,700


Figure 6b. Type-2 cylindrical inclu-
sions (pinwheels with more or less
curved long arms and laminated
aggregates) induced by Bean Yellow
Mosaic Potyvirus in Viciafaba. X
B .-- 27,700











Figure 7. Type-2 cylindrical inclu-
sions (pinwheels and laminated
aggregates induced by Tobacco Etch
Potyvirus in Nicotiana tabacum.
i~'. ^: Tissues were treated with vacuum
S dehydration and OsO4 vapor fixation.
'f1 ~J Note striations on cylindrical inclu-
sion components. Note scattered,
S '" cross-sectioned virus particles (ar-
rows). X 126,200






















F ~ "Yr -" j% c-'


.. .' '* '
S & ,,, '1 '
:,' i .v _, ,. "-*" "" .. .. .r
?,p


jvr, '..


. ,:..


N ~
4' 1 c


Figure 8a. Type-4 cylindrical inclu-
sions (pinwheels, scrolls and short
curved laminated aggregates) in-
duced by Potato Virus Y Potyvirus in
Nicotiana tabacum. X 47,800









Figure 8b. Type-4 cylindrical inclu-
sions (pinwheels, scrolls and short
curved laminated aggregates) in-
duced by Turnip Mosaic
(T isolate) Potyvirus in Brassica
perviridis. X 73,500


Figure 9. Type-2 cylindrical inclu-
sions induced by Tobacco Etch
Potyvirus in Petunia hybrida. Note
layered structure of the laminated
aggregates. Note "eroded" regions in
the upper-left laminated aggregate.
X 82,600


h.


~~ t






















0' **>"


:`"Rf~F~8~~ I~ % W


., V *.


p.~j


.- A y .
:4
Bd


'A,

'*i ..

Li


2w


'.,
..+ .b Ar


Figure 10a. Laminated aggregates
having no apparent associations with
the central portions of cylindrical
inclusions, induced by Tobacco Etch
Potyvirus in Petunia hybrida. Note the
association of one laminated aggre-
gate with endoplasmic reticulum. X
82,600








Figure 10b. Laminated aggregates
having no apparent associations
with the central portions of cylindri-
cal inclusions, induced by Bidens
Mottle Potyvirus in Zinnia elegans.
Note association of laminated
aggregates with the endoplasmic
reticulum. X 47,800
















Figure 11. Laminated aggregates
(tangentially-sectioned) induced by
Tobacco Etch Potyvirus in Petunia
hybrida. Note the absence of layering
in the laminated aggregates. X 47,800


i "'

,B,
ei
4i'

~es~r


~
r























Figure 12. Portion of a cylindrical
inclusion extracted from Potato
Virus Y Potyvirus infected Nicotiana
tabacum. Note striated appearance of
the inclusion's components. X 102,400


2 A'


I-.


I
:rjrsr

~tF~


.i/./jr*I











Figure 13a. Pinwheels induced by
Nothoscordum Mosaic Potyvirus in
Nothoscordum fragrans; vacuum
dehydration, OsO4 vapor fixation.
Inclusions in reversed contrast
exhibit striations. X 140,000




Figure 13b. Pinwheels and scrolls
induced by Wheat Streak Mosaic
Rymovirus in Triticum aestivum;
vacuum dehydration, OsO4 vapor
fixation. Inclusions in reversed
contrast exhibit striations. X 82,600
















Figure 14. Pinwheels and scrolls
induced by Desmodium Mosaic
Potyvirus in Desmodium canum;
conventional dehydration (ethanol
series) and fixation (glutaraldehyde-
OsO4). Note striations on the
inclusions. X 125,000










Figure 15a. Type-1 cylindrical
inclusions (pinwheels and scrolls)
induced by Papaya Ringspot Virus-W
Potvvirus in Cucurbita pepo. X 27,700













Figure 15b. Type-3 cylindrical
inclusions (pinwheels, scrolls and
laminated aggregates) induced by
Turnip Mosaic (Florida isolate)
,j.;, A.: Potyvirus in Brassica perviridis. X 47,800









...

.4;.

4 .;.L~t~iV
f ~r -






.4~



z~ir -~l7~-U



..z ~r.f4,


Figure 16a. Cylindrical inclusions
consisting of pinwheels and scrolls
induced by a Potyvirus in Allium
sativum. X 47,800









Figure 16b. Cylindrical inclusions
consisting of pinwheels, scrolls,
tubes, laminated aggregates and
short curved laminated aggregates
induced by the same Potyvirus
which induced the inclusions in
figure 16a. Tissue samples for 16a
and 16b were taken from the same
Allium sativum leaf. X 27,700













Figure 17. Portions of cylindrical
inclusions consisting of scrolls
sectioned in various planes, induced
by Moroccan Watermelon Mosaic
Potyvirus in Cucurbita pepo. X 16,200


























Figure 18. Type-3 and Type-4
cylindrical inclusions induced by
Watermelon Mosaic Virus-II
Potyvirus in Cucurbita pepo.
Whether this figure represents a
mixture of WMV-II strains or the
result of pleiotropic gene action is
unknown. X 38,500








Table 1. Cytoplasmic Cylindrical Inclusions (CI) Induced by Viruses of the Family Potyviridae, Genus
Potyvirus (Aphid Transmitted).
Viruses Cylindrical Subdivisions Hosts
Inclusions
Type
Alstroemeria Mosaic (S)' EM2-CI

Alstroemeria Streak (TS)3 EM-type 2 (nI)6 Alstroemeria sp.

Amaranthus Leaf Mottle (S) EM-type 1 (I) Nicotiana clevelan

Amazon Lily Mosaic (S) EM-CI -

Aneilema Virus (TS) LM4-type 1 (I) Murdannia nudiflc
EM-type 1 Aneilema aequinoc

Anthoxanthum Mosiac* (TS) EM-type 2 (II) Anthoxanthum od

Aquilegia Virus* (TS) EM-type 2 (II) Aquilegia sp.

Araujia Mosaic (S) EM-type 1 (I) Morrenia odorata

Arracacha Y (TS) No Reports

Artichoke Latent (S) EM-type 2 II Cynara scolymus


Asparagus Virus-1 (S)



Asystasia gangetica Mottle* (TS)

Bean Common Mosaic (S)
Serotype B



BCMV Azuki Bean Mosaic Strain

BCMV Blackeye Cowpea
Mosaic Strain


EM-type 4
EM-type 2


EM-type 2

LM, EM-type 1
EM-type 1



EM-type 1

LM-type 1
LM-type 1
LM, EM-type 1


LM,EM,EMe5-type 1

EM-type 1


Peanut Mild Mottle Strain


EM-type 1

EM-type 1


Refs


dii


ora
tiale

oratum


Datura stramonium
Helianthus annuus
Lathyrus odoratus
Nicotiana benthamiana
N. clevelandii
Phaseolus vulgaris

Chenopodium quinoa
C. amaranticolor
C. quinoa

Nicotiana benthamiana

Phaseolus vulgaris
Phaseolus vulgaris



Phaseolus angularis

Vigna unguiculata
Nicotiana benthamiana
Crotalaria spectabilis
Desmodium canum
Vigna unguiculata
Crotalaria spectabilis
Vigna unguiculata
Crotalaria spectabilis
Desmodium canum
Vigna sesquipedalis
Vigna unguiculata



Nicotiana benthamiana


Refs


465

617

90

574

18
18

154

386

105

76

119,154,303,396,
460,503,524,611
457,458
458
409
174,407
503
458

209
183
250

577,578

421
39,134,152,154,
179,273,351,366,
417,422,423,599

111

163
632
351,627
485
485
165
352, 353
154
154
583
114, 155,355,434,
530,582
383

621








Table 1. (Continued) Cytoplasmic Cylindrical Inclusions (CI) Induced by Viruses of the Family Potyviridae, Genus
Potyvirus (Aphid Transmitted).
Viruses Cylindrical Subdivisions Hosts Refs
Inclusions
Type


Peanut Stripe Strain


Bean Necrosis Mosaic (S)
(Serotype A of BCMV)

Bean Yellow Mosaic (S)


BYMV-Common, Pea Mosaic,
SPMV Strains


EM-type 1
EM-type 1,4
EM-type 4

EM-type 4

EM-type 1


LM-CI


LM,EMe-CI
LM-CIEM-type 2

LM, EM-type 2
LM,EM,EMe-type 2

EM-CI


EM,EMe-type 2

EM-type 2


EM-type 2


Arachis hypogaea
Arachis hypogaea
Arachis hypogaea
Nicotiana benthamiana


Phaseolus vulgaris


Ammi majus
Nicotiana clevelandii
Phaseolus vulgaris
Pisum sativum
Viciafaba
Phaseolus vulgaris
Crocus sativus
Viciafaba
Viciafaba
Lupinus luteus
Pisum sativum
Calanthe sp.
Cucurbita pepo
Proboscidae jussieui
Pisum sativum
Viciafaba
Chenopodium quinoa
Crotalaria sp.
Crotalaria spectabilis
Gibasis geniculata
Gladiolus psittacinus
Iris sp.
Lupinus sp.
Phaseolus vulgaris


Phaseolus vulgaris


28
40
372
28,40,436
5,28,40,511
520
525
523,600
114,289
165
165
265
481
480
261
261
469
587
152
255
626
6
587
89,134,143,154,
297,586


BYMV


Pisum arvense
Pisum sativum

Vicia faba


Beet Mosaic (S)


LM-CI
EM-type 2

EM,EMe-type 2
EM-type 2

EM-type 2


Beta vulgaris
Beta vulgaris

Beta vulgaris
Gomphrena globosa
Dioscorea alata


179
120,202,251,306,
307,570,626
37,39,179,273,
342,344,441,473,
504,570,603,604,
610

40,208
39,148,235,273,
278,393
179
390,391,405,522
475
476


Bidens Mosaic (TS)


EM-type 2








Table 1. (Continued) Cytoplasmic Cylindrical Inclusions (CI) Induced by Viruses of the Family Potyviridae, Genus
Potyvirus (Aphid Transmitted).
Viruses Cylindrical Subdivisions Hosts Refs
Inclusions
Type


Bidens Mottle (S)










Bramble Yellow Mosaic (TS)

Bryonia Mottle (TS)

Canavalia maritima
Mosaic (TS)

Cardamon Mosaic (S)

Carnation Vein Mottle (S)





Carrot Mosaic (TS)


Carrot Thin Leaf (S)

Cassava Brown Streak-Associated (TS)

Cassia Yellow Spot (TS)

Celery Mosaic (S)







Celery Yellow Mosaic (TS)



Chickpea Bushy Dwarf (TS)

Chickpea Filiform (TS)

Chilli Veinal Mottle (S)

Clitoria Yellow
Mosaic (TS)


EMe-type 2

EM-type 2

LM,EMe-type 2

EM,EMe-type 2


LM,EM,EMe-type 2

No Reports

EM-type 1

EM-type 1


EM-type 4

LM-CI
EM-CI
EM-type 2
EM-type 4
EM-type 2,type 4

EM-type 2


No Reports

EM-type 1

EM-type 1

LM-CI
LM,EM-type 1
EM-CI

EM-type 1

EM-type 2

EM-CI
EM-type 1


No Reports

No Reports

EM-CI

LM-CI


Bidens pilosa
Nicotiana edwardsonii
Fittonia verschaffeltii
N. edwardsonii
Lupinus angustifolius
N. edwardsonii
Cichorium endivia
Nicotiana clevelandii
N. edwardsonii
Zinnia elegans



Cucurbita pepo

Canavalia maritima


Elletaria cardamon

Dianthus caryophyllus
Dianthus barbatus
D. barbatus
D. barbatus
D. barbatus

Coriandrum sativum
Daucus carota


Cassia hoffmanseggii

Apium graveolens
Apium graveolens
Ammi majus
Petroselinum sativum
Apium graveolens
Anthriscus cerefolium
Ammi majus


Apium sp.
Coriandrum sativum







Clitoria ternatea


163
232
629
629
164
495
154
117
154
489

168

361

512


198

517
39,502,605
450
154
29

154
135

249

383

447

154
114,488
342
84
3,85,154
83
246

139,140
154
299

191

287

70

349








Table 1. (Continued) Cytoplasmic Cylindrical Inclusions (CI) Induced by Viruses of the Family Potyviridae, Genus
Potyvirus (Aphid Transmitted).


Viruses



Clover Yellow Vein (S)








CYVV Pea Necrosis Strain



Cocksfoot Streak (S)

Colombian Datura (S)

Commelina Mosaic (S)




Cowpea Aphid-Borne Mosaic (S)






Cowpea Green Vein-Banding (S)

Cowpea Rugose Mosaic (TS)

Crinum Mosiac* (TS)

Croatian Clover Mosaic (TS)

Cypripedium calceolus*(TS)

Daphne Virus-Y (TS)


Dasheen Mosaic (S)








Datura Distortion Mosaic (TS)

Datura Mosaic* (TS)

Datura Necrosis (TS)


Cylindrical
Inclusions
Type

LM-CI
LM-type 2
EM-type 2






LM-CI
LM-type 2
EM-type 2

EM-type 4

LM,EM-type 2


LM,EM,EMe-type 2
LM-type 2
EM-type 2

LM-CI

EM-type 1




No Reports

No Reports

EM-type 1

EM-type 2

EM-CI

EM,EMe-type 1


EMe-CI
LM, EMe-CI
LM,EM-type 3
LM,EM,EMe-type 3
EM-type 3




EM-type 3

No Reports

No Reports


Subdivisions Hosts Refs


Nicotiana clevelandi
N. benthamiana
Chenopodium quinoa
Coriandrum sativum
Impatiens sultani
N. clevelandii
Phaseolus vulgaris
Psophocarpus tetragonolobus

Pisum sativum
N. clevelandii
Pisum sativum

Dactylis glomerata

Nicotiana occidental


Commelina diffusa
Rhoeo spathacea
Commelina diffusa

Petunia sp.
Pisum sativum
Crotalaria spectabilis
Phaseolus vulgaris
Vigna unguiculata






Crinum sp.

Nicotiana clevelandii

Cypripedium calceolus

Chenopodium quinoa
Daphne odora

Philodendron selloum
Colocasia antiquorum
Colocasia esculenta
Zantedeschia elliotiana
Alocasia macrorhiza
Colocasia esculenta
Dieffenbachia oerstedii
Vanilla tahitensis


154
159
334
544
118
154,477
267
178

28,47,48
46
46

97

158


424
18
408

41
41
351
261
30,273,351
385

354

364

453

571

345

177
177

195
1
631
630
205
154
205
616

383

379

77








Table 1. (Continued) Cytoplasmic Cylindrical Inclusions (CI) Induced by Viruses of the Family Potyviridae, Genus
Potyvirus (Aphid Transmitted).


Viruses


Datura 437 (TS)


Datura Shoestring (S)

Dendrobium Mosaic (S)

Desmodium Mosaic (TS)


Dioscorea trifida (TS)

Diplandenia Mosaic (TS)

Dock Mottling Mosaic (TS)

Eggplant Green Mosaic (TS)

Eggplant Severe Mottle (TS)


Euphorbia Ringspot (TS)

Ficus carica (TS)

Freesia Mosaic (TS)

Garlic Mosaic (TS)


Garlic Yellow Streak (TS)



Gloriosa Stripe Mosaic (S)

Groundnut Eyespot (S)

Guar Symptomless*(TS)


Guineagrass Mosaic (S)



Habenaria Mosaic (TS)

Helenium Virus Y (S)

Henbane Mosaic (S)


Cylindrical
Inclusions
Type

EM-CI
EM-type 4

EM-type 3

EM-CI

EM-type 1


EM,EMe-type2

EM-type 4

EM-type 2

EM-CI

EM-type 4


EM-type 2

EM-type 4

No Reports

EMe-CI
EM-type 1, 4

EM-CI
EMe-type 1
EM-type 1

EM-type 1

EM-type 4

EM-type 1


EM-type 1



EM-type 1

EM-type 2

LM-CI


EM-CI


Subdivisions Hosts Refs


Datura candida
Datura sp.

Datura metel

Dendrobium sp.

Cyamopsis tetragonoloba
Desmodium canum

Dioscorea trifida

Nicotiana benthamiana

Rumex obtusifolius



Nicotiana clevelandii
N. megalosiphon

Euphorbia milii X lophogona

Ficus carica



Allium sativum
Allium sativum

Allium sativum
Allium sativum
Allium sativum

Gloriosa rothschildiana

Arachis hypogaea

Chenopodium amaranticolor
Cyamopsis tetragonoloba

Brachiaria brizantha
Pennisetum americanum
Zea mays

Habenaria radiata

Chenopodium quinoa

Hyoscyamus niger
Lycopersicon esculentum
Nicotiana tabacum
Datura stramonium
Nicotiana tabacum


125
161

606

262,273

166
154, 158,166

406

451

285

80

321
321

34

203

158

104
104

568
568
459,462

308

145,147,161

216
216

425
315
158

263

317

211
211
211
154
154
2








Table 1. (Continued) Cytoplasmic Cylindrical Inclusions (CI) Induced by Viruses of the Family Potyviridae, Genus
Potyvirus (Aphid Transmitted).
Viruses Cylindrical Subdivisions Hosts Refs
Inclusions
Type


EM,EMe-CI
LM-type 2
LM-type 3
EM-type 3


Hippeastrum Mosaic (S)


Holcus Streak* (TS)


LM-CI


LM, EM-CI
EM-CI
EM-type 2



EM-type 4


Hungarian Datura innoxia Mosaic* (TS) LM-CI
EM-type 4


Hyacinth Mosaic* (TS)

Indian Pepper Mottle (TS)

Iris fulva Mosaic (S)


Iris Mild Mosaic (S)


Iris Severe Mosaic (S)
(Synonym: Bearded Iris Mosaic)


Isachne Mosaic* (TS)

Johnsongrass Mosaic (S)



Kennedia Virus Y (TS)

Konjac Mosaic (S)

Leek Yellow Stripe (S)


Lettuce Mosaic (S)


N. tabacum
N. tabacum
N. tabacum
Datura stramonium
N. tabacum

Hippeastrum equestre
H. hybridum
H. hybridum
H. hybridum
Eucharis grandiflora
Hippeastrum sp.


Holcus mollis

Nicotiana tabacum
Datura innoxia
D. stramonium


No Reports

No Reports

EM-type 4


EMe-CI
EM-type 2
EM,EMe-type 2


EM-CI


EM-type 2
EM,EMe-type 2


EM-type 2

EM-CI
EM-type 1


EM-CI


No Reports

LM-CI
EM-type 3
EM-type 4

EM-type 4,3

No Reports
LM-CI
EM-CI


IV & III


Belamcandia chinensis

Nicotiana clevelandii
Chenopodium quinoa
Iris sp.

Belamcandia chinensis
Chenopodium quinoa
B. chinensis
C. quinoa
Iris sp.

Isachne globosa

Sorghum bicolor
Sorghum halepense
Zea mays

Kennedia rubicunda



Allium porrum
A. porrum
Chenopodium quinoa
A. porrum
A. porrum


217
515
114
248,300
154,158

240
58
66
91
280
445
555

96

464
464
464

158

381

20,22,23


25
154
24, 79,154,357
269
269

580

417
154
187,348,417,463


124

541

45,49
245
443
244,449,597
245


Pisum sativum
P. sativum








Table 1. (Continued) Cytoplasmic Cylindrical Inclusions (CI) Induced by Viruses of the Family Potyviridae, Genus
Potyvirus (Aphid Transmitted).


Viruses


Lily Mild Mottle (S)

Maclura Mosaic
unassigned to a genus




Maize Dwarf Mosaic (S)

MDMV Strain A



MDMV Strain B







MDMV Strain D

MDMV Strain E

MDMV Strain Fleck-20

MDMV Strain G

MDMV Strain 481

Malva Vein Clearing (TS)

Marigold Mottle (TS)

Melilotus Mosaic (TS)

Melon Vein-Banding Mosaic (TS)

Moroccan Watermelon Mosaic (TS)

Mungbean Mosaic* (TS)

Mungbean Mottle (TS)


Cylindrical
Inclusions
Type

EM,EMe-type 2
EM-type 2








EMe-CI

LM-CI

EM-type 2



EM-type 3

EM-CI
EM-type 3
EM-type 1

EM-CI

EM-type 1
EM-type 2
EM-type 3



EM-type 1

EM-type 1

EM-type 1

EM-CI

EM-type 1

EM-type 1

No Reports

No Reports

EM-type 4

EM,EMe-type 1

No Reports

EM-CI


Subdivisions Hosts Refs


P. sativum
Chenopodium amaranticolor
C. quinoa
Lactuca sativa

Picris echioides
Pisum sativum
Spinacia oleracea


Tetragonia expansa

Maclura pomifera
Tetragonia expansa
Maclura pomifera
Nicotiana clevelandii
Tetragonia expansa

Zea mays

Zea mays
Zea mays
Zea mays

Sorghum bicolor
Zea mays
Zea mays
Zea mays
Triticum aestivum
Zea mays


Zea mays

Zea mays

Zea mays

Pennisetumflaccidum

Zea mays

Malva parviflora





Cucurbita pepo

Cucurbita pepo



Vigna radiata


314
126
88, 196,452,531
152,154,257,
273,527
466
417,638
479
155

136,377

468
467,468
468
309,471
467,468

142

312,551
312,419
154,175,348

540
57
175
55
324
154,189,328,
330,348,419

154

154

154

539

348

154,389,394

158

158

252

589

158

567








Table 1. (Continued) Cytoplasmic Cylindrical Inclusions (CI) Induced by Viruses of the Family Potyviridae, Genus
Potyvirus (Aphid Transmitted).
Viruses Cylindrical Subdivisions Hosts Refs


Narcissus Degeneration(S)

Narcissus Latent
unassigned to a genus

Narcissus Late Season Yellows (TS)


Narcissus Yellow Stripe(S)



Nasturtium Mosaic (TS)

Nerine Virus* (TS)




Nothoscordum Mosaic (S)


Onion Yellow Dwarf (S)




Omithogalum Mosaic (S)

Palm Mosaic* (TS)

Papaya Leaf Distortion Mosaic (TS)

Papaya Ringspot (S)
(P isolates)


Papaya Ringspot (S)
(W isolates)






Parsnip Mosaic (S)


Passionfruit Mottle (TS)

Passionfruit Ringspot(TS)


Inclusions
Type

No Reports

EMe-CI
EM-type 2

EM-type 1


LM-type 2
EM-type 2


No Reports

No Reports
EM-type 1



LM-type 1
EM-type 1

LM-CI
LM-type 1
EM-type 1
EM-type 2

No Reports

EM-type 2

EM-type 3


LM-type 1

EM,EMe-type 1
EM-type 1


(II)


(I)


II






(I)



I


I &II






(II)

III

I


LM-CI
LM-type 1
EM,EMe-type 1
EM-type 1

EM-type 1

EM-CI
EM-type 2

LM-type 1

EM-type 1


Nicotiana clevelandii
N. clevelandii

Alstroemeria sp.
Narcissus sp.

Narcissus sp.
Narcissus sp.
Narcissus tazetta




Nerine sp.



Nothoscordumfragrans
N.fragrans

Allium cepa
Allium cepa
A. cepa
A. cepa



Washingtonia robusta

Carica papaya


Carica papaya
Chenopodium amaranticolor
Carica papaya

Carica papaya


Cucurbita pepo
C. pepo
C. pepo
C. pepo

Cucumis sativus

Coriandrum sativum
C. sativum

Passiflora edulis

Cucurbita pepo


562

430
430

75
75

121
121,122,429
154

313

67
G. Balasingham
personal
communication

158
154

43,44
154,371
154
569

141

404

292


113
115
107,154,482
383
230, 304, 564,
613, 628


138,163,392
114,496,498,526
483
129,158,378,498,
526,594
179,204,501

432
433

99

154, 158








Table 1. (Continued) Cytoplasmic Cylindrical Inclusions (CI) Induced by Viruses of the Family Potyviridae, Genus
Potyvirus (Aphid Transmitted).
Viruses Cylindrical Subdivisions Hosts Refs
Inclusions
Type


Passionfruit Woodiness(S)


EM-CI
EM,EMe-type 1
EM-type 1


IV, III


EM,EMe-types 1,2,4
EM-types 1, 4
EM-type 3
EM-type 4


Patchouli Mottle


EM-CI


Peanut Green Mosaic (TS)


Peanut Mosaic (TS)

Peanut Mottle (S)



(Inouye's Isolate)
(Kuhn's Isolate)

Pea Seed-Borne Mosaic (S)


EM-type 1


EM-CI

EM-CI
EM-type 2


EM-type 3
LM-type 3


EM-CI
LM-CI
EM-type 1
EM,EMe-type 1
LM-type 2
EM-type 2
EM-type 4
EM-typel,2,3,4


(I)




II, III


Phaseolus vulgaris
P. vulgaris
P. vulgaris
Passiflora edulis
Phaseolus vulgaris
Nicotiana benthamiana
Phaseolus vulgaris
P vulgaris
Passiflora edulis

Pogostemon patchouli

Arachis hypogaea
Nicotiana benthamiana

Nicotiana glutinosa

Arachis hypogaea
A. hypogaea

Pisum sativum
Viciafaba
Arachis hypogaea


Pisum sativum
Viciafaba
Vfaba
Vfaba
P. sativum
P. sativum
V.faba
P. sativum


Pecteilis Mosaic (TS)

Pepper Mild Mosaic (TS)


Pepper Mottle (S)


Pepper Severe Mosaic (S)


Pepper Veinal Mottle (S)


No Reports

EM-type 2


LM-CI

EM-CI

EMe-CI


EMe-type 1
EM-type 4

LM-type 4

EMe-CI
EM-type 3


Capsicum annuum
Nicotiana glutinosa

Nicotiana tabacum

Capsicum frutescens
N. tabacum
C. annuum
N. edwardsonii
N. tabacum
N. tabacum
C. annuum
N. tabacum
N. tabacum

Nicotiana tabacum
N. tabacum

no inclusions
Lycopersicon esculentum
Nicotiana tabacum


LM
EM-CI


93,341,455
93
417, 572
301
573
356
154
454
99

276a,437a

554
554

621

619
112, 154, 228,
529,620
229,566
275
114


87,214
411
411
272,273
100
110
154
215


133
318

52, 114,138,
154,163
310
439
438
494
494
232
154
114
158,159

171
171








Table 1. (Continued) Cytoplasmic Cylindrical Inclusions (CI) Induced by Viruses of the Family Potyviridae, Genus
Potyvirus (Aphid Transmitted).
Viruses Cylindrical Subdivisions Hosts Refs
Inclusions
Type


EMe-CI,EM-type 4


Petunia hybrida


Perilla Mottle (TS)


Plantain Virus-7 (TS)

Pleioblastus Mosaic (TS)


Plum Pox (S)


Pokeweed Mosaic (S)


Populus Virus* (TS)

Potato Virus A (S)


Potato Virus V (S)

Potato Virus Y (S)


EMe-type 1
EM-type 3

EM-type 1

No Reports

LM-CI
EMe-CI
EMe-type 2
EM-type 2


EMe-type 2
EM-type 3

EM-type 1


EM-CI
LM-CI,EM-type 4


EM-type 2


EM-CI


EM-CI
EM-CI,type 4
EMe-CI
EM-type 1


EM,EMe-type 1
LM-CI,EM-type 1
EM-type 1,4
LM-CI,EM-type 4
EM-type 2
LM-CI,EM-type 2
EMe-CI,EM-type 2
EM-type 4


Perilla frutescens
P.frutescens

Nicotiana benthamiana


Prunus persica
Nicotiana clevelandii
Chenopodiumfoetidum
N. clevelandii

Prunus persica

Phytolacca americana
P. americana



Nicotiana tabacum
N. clevelandii

Nicotiana occidentalis

Capsicum annuum
Datura metel
Nicotiana benthamiana
N. glutinosa
Nicotiana tabacum
Nicotiana tabacum
Capsicum annuum
N. clevelandii
N. glutinosa
N. tabacum
N. tabacum
N. tabacum
N. tabacum
N. tabacum
N. tabacum
N. tabacum
N. tabacum
C. annuum
Datura metel
N. glutinosa
N. tabacum


591,592
51
588
50
313,374,375,
376,397,470,588
169,387


163
154,184,295


181,283


491
35,36
332
10,11
12,152,197,200
473
389
417
179
302,367
233
109
342
154
131,609
114
137
207,565
38, 39,128
545,546,547
158,403
155


PVY Canadian Strain

PVY Australian Strain

Primula Mosaic (TS)


EM-type 4

EM-type 1


LM,EM-type2


N. tabacum

N. clevelandii

Primula obconica








Table 1. (Continued) Cytoplasmic Cylindrical Inclusions (CI) Induced by Viruses of the Family Potyviridae, Genus
Potyvirus (Aphid Transmitted).


Viruses Cylindrical
Inclusions
Type

Primula Mottle (TS) No Reports

Ranunculus Mottle (TS) LM-CI,EM-type2

Reed Canary Mosaic (TS) EM-type 4

Rembrandt Tulip Breaking (S) No Reports

Sorghum Mosaic (S) EM-type 1
(Sorghum Mosaic also contains
Sugarcane Mosaic Virus Strains H and I)

Sugarcane Mosaic-H EM-CI
EM-type 1

EM-type 3
EM-type 4

SCMV-I EM-type 1

South African Passiflora*(S) No Reports

Soybean Mosaic (S) LM-CI
EM-CI

LM-CI,EM-type 1
EM-type 1


Soybean Mosaic Strain A
Strains B, C, D
Strain E

Statice Virus Y (S)

Sugarcane Mosaic (S)

SCMV Strain A


SCMV Strain BC

SCMV Strain ChMV

SCMV Strain D

SCMV Strain E


EMe-type 1,EM-type 3
EM-type 3

LM-type 3
EM-type 1, 4
EM-type 4

EM-type 3
EM-type 4
LM-CI

EM-type 2



EM-type 2
EM-type 3

EM-type 3

EM-CI,type 1,2

EM-type 3

LM-CI
EM-type 4
EM-type 4


Subdivisions Hosts Refs


Ranunculus asiaticus

Phalaris arundinacea


I,IILIV


Zea mays


Zea mays
Zea mays
Sorghum bicolor
Zea mays
Zea mays


I, III, IV






















II
III

III

III


IV


Glycine max
Dolichos lablab
G. max
G. max
G. max


G. max
G. max
Phaseolus vulgaris
Nicotiana benthamiana
G. max
Lupinus albus

G. max
G. max
G. max

Chenopodium quinoa



Zea mays
Zea mays

Zea mays



Zea mays

Zea mays
Zea mays
Sorghum bicolor


380

167

580

136

348



312
175
194,221
154
348

221

56

42, 114,261
108,635
552,585
338
149,175,253,254,
294,339, 342,350,
584,634,637
261
154,273
442
159
256,440
602

581
581
581

342,346



152
154

348

580

154,348

114,528
528
221








Table 1. (Continued) Cytoplasmic Cylindrical Inclusions (CI) Induced by Viruses of the Family Potyviridae, Genus
Potyvirus (Aphid Transmitted).
Viruses Cylindrical Subdivisions Hosts Refs
Inclusions
Type


SCMV Strain M

SCMV Strain SABI

SCMV-Sorghum Red Stripe

SCMV Strain 552

Sunflower Mosaic* (TS)

Sweet Potato Feathery
Mottle(S)=(SP Chlorotic
Leafspot, SP Internal
Cork, SP Russet Crack,
SP Virus A)


Sweet Potato Latent (TS)

Sweet Potato Mild Mottle
unassigned to a genus

Sweet Potato Vein Mosaic (TS)

Sweet Potato Yellow Dwarf
unassigned to a genus

Swordbean Distortion Mosaic (TS)

Tamarillo Mosaic (S)

Teasel Mosaic (TS)

Telfairia Mosaic (S)

Tobacco Etch (S)


EMe-CI

EM-type 3

EM-type 1

EM-type 3

EM-type 4


EM-CI

EM-CI,EM-type 4
EM-type 4


EM-CI

EM-CI


Zea mays


Zea mays

Helianthus annuus


Ipomoea batatas
I. setosa
I. batatas
I. batatas
I. setosa



I. batatas


I. batatas






Nicotiana clevelandii


EM-type 4

EM-CI


EM-type 1

EM-CI,EMe-type 2

No Reports

EM-type 1

LM-type 2

LM-type 2,EMe-CI
LM,EM-type 2
LM,EM,EMe-type 2
EMe-type 2

EM-type 2


EM-type 2


EM-type 2





EM,EMe-type 2


Nicotiana benthamiana

Capsicum chinense
Nicotiana tabacum
N. tabacum
N. tabacum
N. tabacum
Capsicum annuum
N. tabacum
C. annuum
C.frutescens
Cassia tora
Chenopodium amaranticolor
Datura stramonium

Lycopersicon esculentum
Nicotiana benthamiana
N. tabacum




N. tabacum


311

348

383

348

14


444
212, 276, 333
444
333
276

81

237


444

71


383

427

158,563

542

116
27, 154,305,532
515
114, 156,487,538
162
491
234
207
310
484
484
26, 154, 219,
367,400,401
420
326, 332
9,26, 152, 153,
154, 163, 179, 182,
227, 238,484, 518,
534, 608
536, 537
233








Table 1. (Continued) Cytoplasmic Cylindrical Inclusions (CI) Induced by Viruses of the Family Potyviridae, Genus
Pot virus (A hid Transmitted)


Viruses Cylindrical Subdivisions Hosts Refs
Inclusions
Type


Tobacco Vein Banding
Mosaic (TS)

Tobacco Vein Mottling (S)


Tobacco Wilt (TS)

Tomato (Peru) Mosaic (S)


Tongan Vanilla (TS)

Tradescantia & Zebrina(TS)


Trichosanthes Mottle (TS)

Tropaeolum tuberosum Virus-1 (TS)

Tropaeolum tuberosum Virus-2 (TS)

Tulip Band Breaking (S)

Tulip Breaking (S)


Tulip Chlorotic Blotch(S)


Turnip Mosaic (S)


III, IV


EM-type 3,4

EM-CI
EM-type 1

No Reports


EM-type 1
LM-CI,EM-type 4
EM-type 4


EM-type 1

LM-CI
EM-type 1

EMe-CI

No Reports

No Reports

No Reports

LM-CI
EM-CI
EM-type 1


EMe-CI

LM-CI


III, IV


LM-CI


EM-CI
EM,EMe-CI
EM-type 1

LM-CI, type 2

LM-type 3
EM-type 3


Nicotiana tabacum

Nicotiana tabacum
N. tabacum



Nicotiana tabacum
N. occidentalis
N. tabacum

Nicotiana benthamiana

Tradescantia albiflora
T. albiflora

Trichosanthes rostrata


Tulipa sp.
Tulipa sp.
Chenopodium amaranticolor
Liliumformosianum
Lilium sp.

Chenopodium amaranticolor


Alliaria officinalis
Anemone coronaria
Brassica oleraceae
B. rapa
Gomphrena globosa
Matthiola incana
Nicotiana glutinosa
Sinapis alba
Brassica pekinensis
Cichorium endivia
N. glutinosa
Pisum sativum
B. chinensis
B. napus
B. perviridis
B. perviridis

B. rapa
Chenopodium quinoa
Hesperis matronalis
N. clevelandii
Petunia hybrida


509

199,435
8,154,513


456

18
360,362

624

72

73

16,136

372
595
7
622
33


558
236
516
286
410
558
63
412
633
596
64
637
515
515
170
154,163,288,
289,527
13,220,557
297
176
345
220








Table 1. (Continued) Cytoplasmic Cylindrical Inclusions (CI) Induced by Viruses of the Family Potyviridae, Genus
Potyvirus (Aphid Transmitted).
Viruses Cylindrical Subdivisions Hosts Refs
Inclusions
Type


TuMV Strain D

TuMV Strain O

TuMV Strain R


TuMV Florida Isolate

TuMV Tigrida Isolate

Ullucus Mosaic (TS)

Vallota Mosaic (TS)

Watermelon Mosaic Virus-2 (S)


LM,EM,EMe-type 3
EMe-type 3
EMe-type 1, and EM-type 3
EMe-type 1, EM-type 2,3
LM-CI,EM-type 3,4
EMe-type 1,EM-CI, type 4
LM-type 2, EM-type 4
EM-type 1, 4
EM-type 4

EM,EMe-type 3

EM-type 4

EM-type 4
EM,EMe-type 1, 4

LM,EM-type 3

EM-type 4

No Reports

EMe-CI,EM-type 1 (I)

III, IV
EM-CI
EMe-CI
EMe-CI,EM-type 1
EM-type 1


EM-type 3



EMe-CI, EM-type 4
EM-type 4


WMV-2 Florida Isolate

White Bryony Mosaic (TS)

Wild Potato Mosaic (TS)

Wisteria Vein Mosaic (S)




Yam Mosaic (S) (Synonym:


EM-type 3

EM-type 1

EM-type 2

LM-CI
EM-CI, type 1
EM-type 1,4
EM-type 4

EM-type 2


Raphanus sativus

B. perviridis
B. perviridis
B. rapa
N. clevelandii
B. rapa
N. clevelandii
B. oleraceae
Nicotiana glutinosa
Petunia hybrida

Brassica perviridis

N. clevelandii

B. perviridis
B. perviridis

B. perviridis

N. clevelandii


Vallota speciosa


Cucumis sativus
Cucurbita pepo
C. pepo
Chenopodium amaranticolor
Cucumis sativus
Cucurbita pepo

Cucumis sativus
Lupinus angustifolius
Viciafaba

Sesamum indicum
Cucurbita pepo
Lupinus luteus


Cucurbita pepo



Nicotiana clevelandii

Pisum sativum
P. sativum
Pisum sativum
Wisteriafloribunda

Tamus communis


106,273
473
160
232
260
268
559
265
130
60
225

369

342

114,367
369

114

68,154

82

264


473
15
461
17
179
94,290
144
204
114
273
279
102
152, 154
594
492

155

416

158

59
61
62
161

598








Table 1. (Continued) Cytoplasmic Cylindrical Inclusions (CI) Induced by Viruses of the Family Potyviridae, Genus
P=h nrm i Ahid Tran.rmitl-e


Viruses Cylindrical Subdivisions Hosts Refs
Inclusions
Type


Dioscorea Green Banding)

Zoysia Mosaic (TS)

Zucchini Yellow Fleck (S)


Zucchini Yellow Mosaic(S)


Florida Mild Isolate

Reunion Isolate

Type Isolate

Florida Isolate # 2

Florida Isolate # 3


No Reports


LM,EM-type 1
EM-type 1

LM-CI
EMe-CI
EM-CI
LM-CI,EM-type 1
EM-type 1

EMe-type 1
LM,EMe-type 1
EM,EMe-type 1
EM-type 4


I,III,IV


EM-type 1

EM-type 1

EM-type 1

EM-type 3,4

EM-type 3,4


Cucurbita pepo
C. pepo

Cucumis melo
C. sativus
C. sativus
Cucurbita pepo
Cucumis sativus
Cucurbita pepo
Cucurbita pepo
Cucumis sativus

Cucurbita pepo

Cucurbita pepo

Cucurbita pepo

Cucurbita pepo

Cucurbita pepo

Cucurbita pepo


1. (S) = Viruses assigned to the Potyviridae as species by the International Committee On Taxonomy Of Viruses (Barnett, 1991)
2. EM = Electron microscope studies of thin-sections of Potyviridae virus-infected tissues
3. (TS) = Viruses tentatively assigned to the Potyviridae as species (Barnett, 1991)
4. LM = Light microscopy of Potyviridae virus-infected tissues
5. EMe = Electron microscope studies of extracts of plants infected with viruses in the Potyviridae
6. (I), (II) = Viruses assigned to Subdivisions as tentative members because cytological information is limited
= Viruses which have been assigned to the genus Potyvirus, but aphid-transmission has not been demonstrated


601
395

335
365
474
290
347,473
358,359
492
623
209
163

464a

464a

464a

464a


464a








Table 2. Cytoplasmic Cylindrical Inclusions (CI) Induced by Viruses of the Family Potyviridae, Genus Rymovirus
(Eriophyid Mite-Transmission).
Viruses Cylindrical Subdivisions Hosts Refs
Inclusions
Type


Agropyron Mosaic (S)1



Brome Streak Mosaic (TS)3

Hordeum Mosaic (S)

Oat Necrotic Mottle (S)

Ryegrass Mosaic (S)


RMV LM 1 Isolate

RMV LM 3 Isolate

RMV AS Isolate


Wheat Streak Mosaic (S)


EM 2-CI
EM-type 1


EM-type 2

EM-type 1

EM-type 1

EM-CI

EM-type 2


I



II

I

I

II, III


Triticum aestivum
T. aestivum


Hordeum vulgare

Triticum aestivum

Avena sativa

Lolium multiflorum


Avena sativa
L. multiflorum

L. perenne

L. multiflorum

L. multiflorum


EM-type 3


LM4 CI
EM-CI
EM-CI,type 1
EM,EMe-type 1
EM-type 1

EM-type 2
EM-type 3
EM-CI,type 3,4
EM-type 3,4
EM-type 4


IV, III


Polypogon monspeliensis

Triticum aestivum
T. aestivum
T. aestivum
T. aestivum
T. aestivum
Zea mays
Hordeum murinum
T. aestivum
T. aestivum
Zizania aquatica
T. aestivum


325
92,96,154,
324,533,548

413,414

154,324,331

154, 192,193

210,448

472
95, 96, 154,
472,550
150

98


114, 185
53, 325, 535, 561
186,322
323
154,322,324,327
55
500
188
370
32
19,284


Spartina Mottle (TS)


EM-type 2


Spartina angelica


1. (S) = Viruses assigned to the Potyviridae as species by the International Committee On Taxonomy Of Viruses (Barnett, 1991)
2. EM = Electron microscope studies of thin-sections of Potyviridae virus-infected tissues
3. (TS) = Viruses tentatively assigned to the Potyviridae as species (Barnett, 1991)
4. LM = Light microscopy of Potyviridae virus-infected tissues








Table 3. Cytoplasmic Cylindrical Inclusions (CI) Induced by Viruses of the Family Potyviridae, Genus
Bvmovirus (Fungus-Transmitted).


Viruses Cylindrical Subdivisions Hosts Refs
Inclusions
Type


Barley Mild Mosaic (S)1

Barley Yellow Mosaic (S)



Oat Mosaic (S)

Rice Necrosis Mosaic (S)


Wheat Spindle Streak Mosaic (S)


EM2-type 2

EM-CI
EM-type 2
EM-type 2,3


EM-CI
EM-type 2
LM'-CI
EMe', EM-type 2
EM-type 2


EM-CI


EMe-CI
EM-CI, type 2
EM-type 2

EM-CI,type 1, 4
EM-CI,type 4


WSSMV (Synomym: Wheat
Yellow Mosaic Virus)


LM-CI
EM-type 2


Hordeum vulgare

Hordeum vulgare
H. vulgare
H. vulgare

Avena sativa
A. sativa
Bracharia ramosa
Oryza sativa
0. sativa

Triticum aestivum


T. aestivum
T. aestivum
T. aestivum
T. duram
T. aestivum
T. aestivum

T. aestivum
T. aestivum


= Viruses assigned to the Potyviridae as species by the International Committee On Taxonomy Of Viruses (Barnett, 1991)
= Electron microscope studies of thin-sections of Potyviridae virus-infected tissues
= Light microscopy of Potyviridae virus-infected tissues
= Electron microscope studies of extracts of plants infected with viruses in the Potyviridae


259

275
151,351a
388

223
154,231
190
271
274,437


54, 241,325,
329,363,549,
613,614
543
322
19,231,277
514
242
154


1.(S)
2. EM
3. LM
4. EMe








Table 4. Viruses Exhibiting Potyvirus Properties But Which Are Presently Unassigned to the Family Potyvirideae.

Viruses Cylindrical Subdivisions Hosts Refs
Inclusions
Type

Amaranthus lividus EM'-type 3 III Amaranthus lividus 521

Arachis Mottle EM-type 4 IV Arachis hypogaea 201

Cassia Severe Mosaic EM-type 1 (I) Nicotiana clevelandii 602a

Chickpea Distortion Mosaic EM-type 1 (I) Cicer arietinum 384

Chickpea Marathwada EM-type 1 (I) Cicer arietinum 382

Crocus Y-C EM-type 2,4 IV Crocus sp. 31

Daphne mezereum EM-type 3 III Daphne mezereum 556

Elephant Grass Mosaic EM-type 1 (I) Pennisetum purpureum 399

Gladiolus Potyvirus EM-type 3 III Viciafaba 293

Kalanchoe Mosaic No Reports Kalanchoe blossfeldiana 258

Sesame Potyvirus LM2-CI IV Sesamum indicum 553
LM-type 4 S. indicum (R.G. Christie


Sesame Yellow Mosaic EM-type 1 (I)

Spiranthes Mosaic EM-type 3 III

Stylosanthes Mild Chlorosis EM-type 2 (II)

Tuberose Mottle EM-type 2 (II)


1. EM = Electron microscope studies of thin-sections of Potyviridae virus-infected tissues
2. LM = Light microscopy of Potyviridae virus-infected tissues


Sesamum indicum

Chenopodium amaranticolor

Stylosanthes sp.

Polianthes tuberosa


unpublished)

542a

101

426

243








Table 5. Viruses Whose Cylindrical Inclusions Have Not Been Characterized.

Viruses Species

Alstroemeria Mosaic S1

Chilli Veinal Mottle S

Clitoria Yellow Mosaic TS2

Cypripedium calceolus TS

Dendrobium Mosaic S

Eggplant Green Mosaic TS

Kennedia Virus Y TS

Lily Mild Mottle S

Mungbean Mottle TS

Patchouli Mottle TS

Peanut Mosaic TS

Pea Seed-Borne Mosaic S

Sweet Potato Latent TS

Sweet Potato Mild Mottle

Sweet Potato Yellow Dwarf

Trichosanthes Mottle TS

Tulilp Chlorotic Blotch S


1. S = Viruses assigned to the Potyviridae as species by the International Committee On Taxonomy Of Viruses (Bamett, 1991)
2. TS = Viruses tentatively assigned to the Potyviridae as species (Barnett, 1991)








Table 6. Viruses Inducing Type-1 Cylindrical Inclusions Constitute Subdivision I.

Viruses Species Subdivision

GENUS POTYVIRUS

Amaranthus Leaf Mottle S1 (I)2

Aneilema Virus TS3 (I)

Araujia Mosaic S (I)

Bean Common Mosaic S I

Bean Necrosis Mosaic S (I)

Bryonia Mottle TS (I)

Canavalia maritima Mosaic TS (I)

Cassava Brown Streak-Associated TS (I)

Cassia Yellow Spot TS (I)

Celery Mosaic S I

Celery Yellow Mosaic TS I

Cowpea Aphid-Borne Mosaic S I

Crinum Mosaic TS (I)

Daphne Virus-Y TS (I)

Desmodium Mosaic TS I

Garlic Yellow Streak TS I

Gloriosa Stripe Mosaic S (I)

Guar Symptomless TS (I)

Guineagrass Mosaic S I

Habenaria Mosaic TS (I)

Johnsongrass Mosaic S I

Maize Dwarf Mosaic S I (also Subdivi

Maize Dwarf Mosaic
(Strains D, E, Fleck-20, 481) (I)
(Strain A) I (also Subdivi

Malva Vein Clearing TS I

Moroccan Watermelon Mosaic TS (I)

Narcissus Late Season Yellows TS (I)

Nerine Virus TS (I)

Nothoscordum Mosaic S I

72


sions III & IV)


sion III)








Table 6. (Continued) Viruses Inducing Type-1 Cylindrical Inclusions Constitute Subdivision I.

Viruses Species Subdivision


Onion Yellow Dwarf

Papaya Ringspot (Strains P & W)

Passionfruit Mottle

Passionfruit Ringspot

Peanut Green Mosaic

Plantain Virus-7

Populus Virus

Potato Virus Y (Australian
Isolate)

Sorghum Mosaic

Sugarcane Mosaic
(Sorghum Red Stripe Strain)

Telfairia Mosaic

Tobacco Vein Mottling

Tongan Vanilla

Tradescantia & Zebrina Virus

Tulip Breaking

Vallota Mosaic

White Bryony Mosaic

Zucchini Yellow Fleck

Zucchini Yellow Mosaic

GENUS RYMOVIRUS

Agropyron Mosaic

Hordeum Mosaic

Oat Necrotic Mottle


also Subdivision IV)


(also Subdivisions III & IV)

(also Subdivisions III & IV)



















(also Subdivisions III, IV)


1. S = Viruses assigned to the Potyviridae as species by the International Committee On Taxonomy Of Viruses (Barnett, 1991)
2. (I) = The virus has been tentatively assigned to Subdivision-I because only limited cytological information is available.
3. TS = Viruses tentatively assigned to the Potyviridae as species (Barnett, 1991)









Table 7. Viruses Inducing Type-2 Cylindrical Inclusions Constitute Subdivision II.

Viruses Species Assignments

GENUS POTYVIRUS

Alstroemeria Streak TS1 (II)2

Anthoxanthum Mosaic TS (II)

Aquilegia Virus TS (II)

Artichoke Latent S3 I

Asystasia gangetica Mottle TS II

Bean Yellow Mosaic S II

Beet Mosaic S II

Bidens Mosaic TS (II)

Bidens Mottle S II

Carrot Mosaic TS II

Clover Yellow Vein S II

Colombian Datura S (II)

Commelina Mosaic S II

Croatian Clover Mosaic TS (II)

Dioscorea trifida TS (II)

Dock Mottling Mosaic TS (II)

Euphorbia Ringspot TS (II)

Helenium Virus Y S (II)

Hippeastrum Mosaic S II

Iris Mild Mosaic S II

Iris Severe Mosaic S II

Isachne Mosaic TS (II)

Lettuce Mosaic S I

Maclura Mosaic unassignedd) II

Narcissus Latent unassignedd) (II)

Narcissus Yellow Stripe S II

Palm Mosaic TS (II)

Parsnip Mosaic S (II)








Table 7. (Continued) Viruses Inducing Type-2 Cylindrical Inclusions Constitute Subdivision II.

Viruses Species Assignments

Peanut Mottle S II (also Subdivision III)

Pepper Mild Mosaic TS (II)

Plum Pox S II

Potato VirusV S II

Primula Mosaic TS (II)

Ranunculus Mottle TS (II)

Statice Virus Y S II

Tamarillo Mosaic S (II)

Tobacco Etch S II

Wild Potato Mosaic TS II

Yam Mosaic S II

GENUS RYMOVIRUS

Ryegrass Mosaic S II (also Subdivision III)

Brome Streak TS II

Spartina Mottle TS (II)

GENUS BYMOVIRUS

Barley Mild Mosaic S (II)

Oat Mosaic S II

Rice Necrosis Mosaic S II


1. TS = Viruses tentatively assigned to the Potyviridae as species (Bamett, 1991)
2. (II) = The virus has been tentatively assigned to Subdivision-II because only limited cytological information is available.
3. S = Viruses assigned to the Potyviridae as species by the International Committee On Taxonomy Of Viruses (Barnett, 1991).








Table 8. Viruses Inducing Type-3 Cylindrical Inclusions Constitute Subdivision III.

Viruses Species Subdivision Assignments

GENUS POTYVIRUS

Dasheen Mosaic S1 III

Datura Shoestring S III

Datura Distortion Mosaic TS2 III

Henbane Mosaic S III

Leek Yellow Stripe S III (also Subdivision IV)

Maize Dwarf Mosaic (strains A,B) S III (also Subdivisions I & IV)

Passionfruit Woodiness S III (also Subdivision IV)

Peanut Mottle (Inouye & Kuhn S III (also Subdivision II)
Isolates)

Pepper Severe Mosaic S III

Perilla Mottle TS III

Pokeweed Mosaic S III

Sorghum Mosaic (Sugarcane Mosaic S III (also Subdivisions I & IV)
Strain H)

Soybean Mosaic (Strain A & other S III (also Subdivision IV)
Isolates)

Sugarcane Mosaic (Strains A, BC, S III (also Subdivisions I & IV)
D, ChMV, 552, SABI)

Turnip Mosaic (Strain D) S III (also Subdivision IV)

(Florida Isolate and others) S III (also Subdivision IV).

Watermelon Mosaic Virus-2 S III (also Subdivision IV)
(Florida Isolate & others)

Zucchini Yellow Mosaic S III (also Subdivisions I & IV)

GENUS RYMOVIRUS

Ryegrass Mosaic (AS Isolate) S III (also Subdivision II)

Wheat Streak Mosaic S III (also Subdivision IV)

GENUS BYMOVIRUS

Barley Yellow Mosaic S III


1. S = Viruses assigned to the Potyviridae as species by the International Committee On Taxonomy Of Viruses (Barnett, 1991)
2. TS = Viruses tentatively assigned to the Potyviridae as species (Barnett, 1991)








Table 9. Viruses Inducing Type-4 Cylindrical Inclusions Constitute Subdivision IV.

Viruses Species Assignments

GENUS POTYVIRUS
Asparagus Virus-1 S1 IV

Bean Common Mosaic (Azuki Bean S IV (also Subdivision I)
Mosaic & Peanut Stripe Strains)

Cardamon Mosaic S IV

Carnation Vein Mottle S IV

Cocksfoot Streak S IV

Datura 437 S IV

Diplandenia Mosaic TS2 IV

Eggplant Severe Mottle TS IV

Ficus carica Virus TS IV

Garlic Mosaic TS IV

Groundnut Eyespot S IV

Holcus Streak TS IV

Hungarian Datura innoxia Mosaic TS IV

Iris fulva Mosaic S IV

Leek Yellow Stripe S IV (also Subdivision III)

Maize Dwarf Mosaic (Strain SP) S IV (also Subdivisions I &

Melon Vein-Banding Mosaic TS IV

Passionfruit Woodiness S IV (also Subdivision III)

Pepper Mottle S IV

Pepper Veinal Mottle S IV

Potato Virus A S IV

Potato Virus Y S IV (also Subdivision I)

Reed Canary Mosaic TS IV

Sorghum Mosaic (Sugarcane Mosaic S IV (also Subdivisions I &
Strain H)

Soybean Mosaic (Strains B, C, D S IV (also Subdivision III)
& others)

Sugarcane Mosaic (Strain E) S IV (also Subdivisions I &

Sunflower Mosaic TS IV


III)


III)


III)








Table 9. (Continued) Viruses Inducing Type-4 Cylindrical Inclusions Constitute Subdivision IV.

Viruses Species Assignments


Sweet Potato Feathery Mottle

Sweet Potato Vein Mosaic

Tomato (Peru) Mosaic

Turnip Mosaic (Tigrida Isolate,
Strain O, R & others)

Watermelon Mosaic Virus-2

Wisteria Vein Mosaic

Zucchini Yellow Mosaic

GENUS RYMOVIRUS

Wheat Streak Mosaic

GENUS BYMOVIRUS

Wheat Spindle-Streak Mosaic


IV

IV

IV

IV (also Subdivision III)


IV (also Subdivision III)

IV

IV (also Subdivisions I & III)



IV (also Subdivision III)


1. S = Viruses assigned to the Potyviridae as species by the International Committee On Taxonomy Of Viruses (Barnett, 1991)
2. TS = Viruses tentatively assigned to the Potyviridae as species (Barnett, 1991)








Table 10. Viruses of the Potyviridae with No Cytological Studies.

Viruses Species

GENUS POTYVIRUS

Arracacha-Y TS

Bramble Yellow Mosaic TS
Carrot Thin Leaf S1
Chickpea Bushy Dwarf TS2
Chickpea Filiform TS
Cowpea Green Vein-Banding S
Cowpea Rugose Mosaic TS
Datura Mosaic TS
Datura Necrosis TS
Freesia Mosaic TS
Hyacinth Mosaic TS
Indian Pepper Mottle TS

Konjac Mosaic S
Marigold Mottle TS
Melilotus Mosaic TS
Mungbean Mosaic TS
Narcissus Degeneration S
Nasturtium Mosaic TS
Ornithogalum Mosaic S
Pecteilis Mosaic TS
Pleioblastus Mosaic TS
Primula Mottle TS
Rembrandt Tulip Breaking S
South African Passiflora TS
Teasel Mosaic TS
Tobacco Wilt TS
Tropaeolum tuberosum Virus-1 TS
Tropaeolum tuberosum Virus-2 TS
Tulip Band Breaking S
Ullucus Mosaic TS
Zoysia Mosaic TS

1. S = Viruses assigned to the Potyviridae as species by the International Committee On Taxonomy Of Viruses (Barnett, 1991)
2. TS = Viruses tentatively assigned to the Potyviridae as species (Barnett, 1991)








































































Florida Agricultural Experiment Station, Institute of Food and Agricultural Sciences, University of Florida, Richard L. Jones, Dean for Research, publishes
this information to further programs and related activities, available to all persons regardless of race, color, age, sex, handicap or national origin. For
information about alternate formats, contact the Educational Media and Services Unit, University of Florida, PO Box 110810, Gainesville, FL 32611-
0810. This information was published May 1996 as Bulletin 894 Florida Agricultural Experiment Station.
ISSN 0096-607X




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