Table Of ContentVIROIDS
and
SATELLITES
Cover photographs
Front cover
Electron micrograph of small circular coconut cadang-cadang viroid
molecules. (Courtesy John W. Randles.)
Coconut cadang-cadang viroid induced non-necrotic yellow spotting of
a coconut palm leaflet (right) compared with an asymptomatic leaflet
(left). (Courtesy John W.Randles.)
Potato spindle tuber viroid affected potato tuber showing typical
spindle shape (right) compared with a normal potato tuber (left).
(Courtesy Theodor O.Diener.)
Extreme leaf chlorosis (peach calico) symptoms induced by certain
variants of peach latent mosaic viroid. (Courtesy Francesco Di Serio and
Ricardo Flores.)
Symptoms of citrus exocortis viroid infection on a citron fruit (right)
compared with anormal fruit (left).(Courtesy Joseph S.Semancik.)
Severe symptoms of a variant of cucumber mosaic virus Tfn-satellite
RNA on tomato fruit cv. Italpeel. (Courtesy Donato Gallitelli and the
late Makis Vovlas.)
Symptoms of the dapple apple variant of apple scar skin viroid on
apple fruit cv.Starkrimson. (Courtesy Jean-ClaudeDesvignes.)
Back cover
Symptoms of chrysanthemum stunt viroid on white chrysanthemum
flower (right) showing reduced size and quality compared with a
normalflower (left). (Courtesy Teruo Sano.)
Avocado sunblotch viroid induced depression and distortion symptoms
on anavocado fruit. (Courtesy David N.Kuhn.)
VIROIDS
and
SATELLITES
Edited by
A H
HMED ADIDI
U.S.DepartmentofAgriculture,Beltsville,MD,UnitedStates
R F
ICARDO LORES
PolytechnicUniversityofValencia-CSIC,Valencia,Spain
J W. R
OHN ANDLES
TheUniversityofAdelaide,WaiteCampus,GlenOsmond,Australia
P P
ETER ALUKAITIS
SeoulWomen’sUniversity,Seoul,SouthKorea
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List of Contributors
Mazen Alazem Institute of Plant and Microbial Biology, Academia Sinica,
Taipei,Taiwan
HaniZ.Al-Tuwariqi MinistryofAgriculture,Riyadh,SaudiArabia
Marina Barba CREA-Research Centre for Plant Protection and Certification,
Rome,Italy
Iraklis N. Boubourakas Directorate of Rural Economy and Veterinary of
Piraeus,AtticaPrefecture,Attica,Greece
KadriyeC¸ag˘layan MustafaKemalUniversity,Antakya,Turkey
ThierryCandresse INRAandUniversityofBordeaux,Bordeaux,France
VicenteConejero PolytechnicUniversityofValencia-CSIC,Valencia,Spain
Elena Dadami RLP AgroScience, AlPlanta-Institute for Plant Research,
Neustadt-Mußbach,Germany
Athanasios Dalakouras RLP AgroScience, AlPlanta-Institute for Plant
Research,Neustadt-Mußbach,Germany
Jose´-AntonioDaro`s PolytechnicUniversityofValencia-CSIC,Valencia,Spain
MarcosDelaPen˜a PolytechnicUniversityofValencia-CSIC,Valencia,Spain
SoniaDelgado PolytechnicUniversityofValencia-CSIC,Valencia,Spain
FrancescoDiSerio NationalResearchCouncil,Bari,Italy
JonathanDixon SeekaKiwifruitIndustriesLimited,TePuke,NewZealand
Nuria Duran-Vila Valencian Institute of Agricutural Research-IVIA, Valencia,
Spain
KhaledA.El-Dougdoug AinShamsUniversity,Cairo,Egypt
AmineElleuch UniversityofSfax,Sfax,Tunisia
Francesco Faggioli CREA-Research Centre for Plant Protection and
Certification,Rome,Italy
RicardoFlores PolytechnicUniversityofValencia-CSIC,Valencia,Spain
Aurora Fraile Center for Plant Biotechnology and Genomics, Polytechnic
UniversityofMadrid-INIA,Madrid,Spain
SelmaGago-Zachert LeibnizInstituteofPlantBiochemistry,Halle,Germany
DonatoGallitelli UniversityofBariAldoMoro,Bari,Italy
Fernando Garcı´a-Arenal Center for Plant Biotechnology and Genomics,
PolytechnicUniversityofMadrid-INIA,Madrid,Spain
xxv
xxvi
LISTOFCONTRIBUTORS
Andrew D.W. Geering The University of Queensland, St Lucia, QLD,
Australia
LucianoGiunchedi UniversityofBologna,Bologna,Italy
GustavoGo´mez PolytechnicUniversityofValencia-CSIC,Valencia,Spain
EbenezerA.Gyamera UniversityofCambridge,Cambridge,UnitedKingdom
Nuredin Habili The University of Adelaide, Waite Campus, Glen Osmond,
SA,Australia
AhmedHadidi U.S.DepartmentofAgriculture,Beltsville,MD,UnitedStates
Vipin Hallan CSIR-Institute ofHimalayan BioresourceTechnology, Palampur,
HimachalPradesh,India
Rosemarie W. Hammond U.S. Department of Agriculture, Beltsville, MD,
UnitedStates
Dagmar Hanold The University of Adelaide, Waite Campus, Glen Osmond,
SA,Australia
TatsujiHataya HokkaidoUniversity,Sapporo,Japan
CarmenHerna´ndez PolytechnicUniversityofValencia-CSIC,Valencia,Spain
NiHong HuazhongAgriculturalUniversity,Wuhan,China
MunetakaHosakawa KyotoUniversity,Kyoto,Japan
Yau-HeiuHsu NationalChungHsingUniversity,Taichung,Taiwan
Chung-ChiHu NationalChungHsingUniversity,Taichung,Taiwan
Ying-WenHuang NationalChungHsingUniversity,Taichung,Taiwan
Delano James Centre for Plant Health-Canadian Food Inspection Agency,
NorthSaanich,BC,Canada
HendryJoseph UniversitiTeknologiMARA(Sabah),Sabah,Malaysia
KritonKalantidis UniversityofCrete,Heraklion,Greece
MariaS.Kaponi Formerly,HirosakiUniversity,Hirosaki,Japan
Mikyeong Kim National Institute of Agricultural Sciences, RDA, Wanju-gun,
SouthKorea
GaryR.Kinard U.S.DepartmentofAgriculture,Beltsville,MD,UnitedStates
LihL.Kong UniversitiPutraMalaysia,Selangor,Malaysia
Natalia Kovalskaya U.S. Department of Agriculture, Beltsville, MD, United
States
DavidN.Kuhn U.S.DepartmentofAgriculture,Miami,FL,UnitedStates
PanayotaE.Kyriakopoulou AgriculturalUniversityofAthens,Athens,Greece
IreneLavagi UniversityofCalifornia,Riverside,CA,UnitedStates
Shi-FangLi ChineseAcademyofAgriculturalSciences,Beijing,China
Na-Sheng Lin Institute of Plant and Microbial Biology, Academia Sinica,
Taipei,Taiwan
Kai-ShuLing U.S.DepartmentofAgriculture,Charleston,SC,UnitedStates
Ming-RuLiou NationalChungHsingUniversity,Taichung,Taiwan
xxvii
LISTOFCONTRIBUTORS
Purificacio´nLiso´n PolytechnicUniversityofValencia-CSIC,Valencia,Spain
Amparo Lo´pez-Carrasco Polytechnic University of Valencia-CSIC, Valencia,
Spain
Marta Luigi CREA-Research Centre for Plant Protection and Certification,
Rome,Italy
A´ngel-Emilio Martı´nez de Alba Polytechnic University of Valencia-CSIC,
Valencia,Spain
TizianaMascia UniversityofBariAldoMoro,Bari,Italy
ChikaraMasuta HokkaidoUniversity,Sapporo,Japan
JaroslavMatousˇek AcademyofSciencesoftheCzechRepublic(ASCR), Cˇeske´
Budeˇjovice,CzechRepublic
Marie-ChristineMaurel SorbonneUniversity,Paris,France
EllisT.M.Meekes Naktuinbouw,Roelofarendsveen,TheNetherlands
SofiaMinoia PolytechnicUniversityofValencia-CSIC,Valencia,Spain
BeatrizNavarro NationalResearchCouncil,Bari,Italy
XianzhouNie AgricultureandAgri-FoodCanada,Fredericton,NB,Canada
Robert A. Owens U.S. Department of Agriculture, Beltsville, MD, United
States
VicentePalla´s PolytechnicUniversityofValencia-CSIC,Valencia,Spain
PeterPalukaitis SeoulWomen’sUniversity,Seoul,SouthKorea
Dattaraj B. Parakh ICAR-National Bureau of Plant Genetic Resources, New
Delhi,India
Jean-PierrePerreault UniversityofSherbrooke,Sherbrooke,QC,Canada
Edward V. Podleckis U.S. Department of Agriculture, Riverdale, MD, United
States
JesseD.Pyle HarvardMedicalSchool,Boston,MA,UnitedStates
John W. Randles The University of Adelaide, Waite Campus, Glen Osmond,
SA,Australia
DetlevRiesner HeinrichHeineUniversity,Du¨sseldorf,Germany
M.JudithB.Rodriguez PhilippineCoconutAuthority,Albay,Philippines
Johanna W. Roenhorst National Plant Protection Organization, Wageningen,
TheNetherlands
Marilyn J. Roossinck Pennsylvania State University, University Park, PA,
UnitedStates
LuisaRubino NationalResearchCouncil,Bari,Italy
Jesus A. Sa´nchez-Navarro Polytechnic University of Valencia-CSIC, Valencia,
Spain
TeruoSano HirosakiUniversity,Hirosaki,Japan
Karen-Beth G. Scholthof Texas A&M University, College Station, TX, United
States
xxviii
LISTOFCONTRIBUTORS
PedroSerra PolytechnicUniversityofValencia-CSIC,Valencia,Spain
HanakoShimura HokkaidoUniversity,Sapporo,Japan
RudraP.Singh AgricultureandAgri-FoodCanada,Fredericton,NB,Canada
DijanaSˇkoric´ UniversityofZagreb, Zagreb,Croatia
GiuseppeStancanelli EuropeanFoodSafetyAuthority,Parma,Italy
GerhardSteger HeinrichHeineUniversityDu¨sseldorf,Germany
Anna Taglienti CREA-Research Centre for Plant Protection and Certification,
Rome,Italy
MatildeTessitori UniversityofCatania,Catania,Italy
SathisS.Thanarajoo UniversitiPutraMalaysia,Selangor,Malaysia
Antonio Tiberini Mediterranea University of Reggio Calabria, Reggio
Calabria,Italy
EnzaM.Torchetti NationalResearchCouncil,Bari,Italy
MinaTsagris UniversityofCrete,Crete,Greece
TaroTsushima HirosakiUniversity,Hirosaki,Japan
GanesanVadamalai UniversitiPutraMalaysia,Selangor,Malaysia
Jacobus Th.J. Verhoeven National Plant Protection Organization,
Wageningen,TheNetherlands
GeorgiosVidalakis UniversityofCalifornia,Riverside,CA,UnitedStates
Qiao-ChunWang NorthwestA&FUniversity,Xianyang,China
Michael Wassenegger RLP AgroScience, AlPlanta-Institute for Plant Research,
Neustadt-Mußbach,Germany
Stephan Winter Leibniz Institute, DSMZ- German Collection of
MicroorganismsandCellCultures,Braunschweig,Germany
Wen-XingXu HuazhongAgriculturalUniversity,Wuhan,China
XiulingYang ChineseAcademyofAgriculturalSciences,Beijing,China
Yongjiang Zhang Chinese Academy of Inspection and Quarantine, Beijing,
China
ZhiboZhang NorthwestA&FUniversity,Xianyang,China
Xueping Zhou Chinese Academy of Agricultural Sciences, Beijing, China;
ZhejiangUniversity,Hangzhou,China
ShuifangZhu ChineseAcademyofInspectionandQuarantine,Beijing,China
Foreword
I am delighted to write a Foreword for this well-conceived book,
which, with the prominent authors chosen, is assured to take its place
as an up-to-date, complete, and dependable textbook on the plant-
infecting subviral viroids and satellites for years to come.
The third majorextensioninthe history ofthe biospherewas ushered
in by the discovery of viroids in 1971, following the earlier discoveries
of the “subvisible” microorganisms by Antonie van Leeuwenhoek in
1675 and of the “submicroscopic” viruses by Dmitri Iosifovich
Ivanovsky in 1892. The status of viroids is recognized by the
International Committee for Virus Taxonomy (ICTV) with the creation
of a new order of subviral agents, which currently includes two families,
eightgenera, and32 species of viroids.
A great interest in viroids among plant virologists occurred immedi-
ately after their discovery and the important biological, biochemical,
and evolutionary significance of subviral agents has been acknowledged
by many other scientists who confirmed the fundamental nature of vir-
oids (and, implicitly, of other subviral agents), which, like viruses, are
replicated in infected cells.
It is now accepted that viroids, like viruses, share the most character-
istic property of living beings: in an appropriate environment, they are
able to generate copies of themselves; in other words, they are endowed
with autonomous replication (and evolution). It is in this framework
that viroids represent the frontier of life (infectious agents consisting of
only 246(cid:1)401 ribonucleotides), an aspect that should attract the atten-
tion of anybody interested in biology.
TheodorO. Diener
xxix
Preface
Many plant diseases currently known to be caused by viroids were
first recognized because of their damaging economic impact many years
ago.PotatospindletuberdiseasewasfirstdescribedintheUnitedStates
in the early 1920s and its infectious agent was identified by Theodor O.
Diener of the US Department of Agriculture, Beltsville, Maryland, in
1971. He described it as a free RNA of 25,000(cid:1)110,000 Daltons, much
smallerthanaviralgenome,andnameditas“viroid”in1972.Similarly,
the “exocortis” disease of citrus was first described in the late 1940s in
the United States and in 1972, Joseph S. Semancik of the University of
California, Riverside, showed that its causal agent is also a viroid of
125,000 Daltons. Cadang-cadang disease of coconut palms in the
Philippines, so named because it leads to the premature decline and
deathoftrees,wasfirstdescribedearlyinthe20thcenturybutitwasnot
until 1975 that John W. Randles of the University of Adelaide, South
Australia,demonstratedthatthediseaseiscausedbyaviroid.By1977,it
was found that viroids cause economically important diseases in
vegetable crops, fruit trees, ornamentals, and palm species. The host
range of viroids has expanded over the years to include grapevine and
other cultivated and wild plant species. Viroid molecules are very small
circular RNAs, ranging from 246 to 401 nucleotides in length and with a
high degree of internal base-pairing. Their discovery has challenged the
central dogma of genetic inheritance because unlike viruses they are
naked and lack protein-coding ability. Although found only in plants,
their discovery paved the way for other subviral agents such as prions
and hepatitis delta virus to be recognized as pathogens of eukaryotic
organisms.ViroidsarethesmallestknowninfectiousRNAs.
Basil Kassanis of the Rothamsted Experiment Station, Harpenden,
Hertfordshire, UK, coined the name “satellite” in 1962 for very small
virus particles he found in some cultures of tobacco necrosis virus
(TNV). He demonstrated that these 17-nm viral satellite particles were
unable to replicate in the absence of the larger 30-nm TNV particles.
Similarly, in 1972 Irving R. Schneider of the US Department of
Agriculture in Beltsville found nonessential but biologically active satel-
lite RNA molecules in preparations of various other viruses. In 1972, a
devastating disease epidemic swept through tomato fields of the French
Alsace province destroying the crop. The causal agent was not revealed
until 1977, when Jacobus M. Kaper of the US Department of Agriculture
xxxi
