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Β© 2018, IRJET | Impact Factor value: 7.211 | ISO 9001:2008 Certified Journal | Page 1865
Research developments in begomovirus in legumes : Past chievements,
Present scenario and Future thrust areas
Lavania Pranesh1, Singh Priya2, Sharma Manju3
1&2Department of Plant Pathology, College of Agriculture, G.B. Pant University of Agriculture and Technology,
Pantnagar, Uttarakhand, India
3Assistant Professor, Department of Plant Pathology, College of Agriculture, G.B. Pant University of Agriculture and
Technology, Pantnagar, Uttarakhand, India
---------------------------------------------------------------------***---------------------------------------------------------------------
Abstract - Begomovirus – A well recognized genus of plant
viruses classified under Geminiviridae family of Group II
(single stranded circular DNA) is known to cause virulence
in a large range of hosts including tomato, okra, cotton,
cassava, bittergourd, chilli, croton, cucumber, eggplant,
jatropha, mentha, mesta, papaya, potato, tobacco, legumes
and many more. It is believed that organism has caused
major economic losses on cropsof all types. However,owing
to today’s demand of increased production, land area and
productivity of pulses for future food and nutritional
security, this paper will summarize collectively the research
study and experiments conducted on legumes till date,
present research work being conducted and future thrust
areas on begomoviruses w.r.t. legumes like common bean
(Phaseolus vulgaris), mung bean (Vigna radiata), urd bean
(Vigna mungo), pigeon pea (Cajanuscajan),mothbean(Vigna
aconitifolia), cowpea (Vignaungiculata),velvetbean(Mucuna
pruriens), frenchbean (Phaseolus vulgaris), soybean (Glysine
max), long yard bean (Vigna sesquipedalis) etc. Prime focus
areas of this review will include molecular characterization,
DNA replication, phylogenetic analysis and infectivity
patterns on host legumes.
Key Words: Begomovirus, legumes, Bemisia tabaci, yellow
mosaic disease, bean common mosaic virus
1. Introduction-
Among all the diseases of plants, viral diseases are the most
complex and the least known. Intensive research in the
recent past has led to showcase some facts with clear
understanding. This review will deal with morphology,
taxonomy, genome, population genomics, gene expression,
economic impact, host range, transmission, molecular
characterization, antigenicity, infectivityandmanagementof
begomoviruses that forms largest group of disease causing
viruses of plants and specifically of legumes.
2. Morphology-
Virions of geminivirus sub group III are germinate .i.e.
twinned with two incomplete icosahedra. They are non-
enveloped in nature and have a common dimensionof30-38
nm * 18-28 nm (length*diameter) with 22 pentameric
capsomers per nucleocaspid and 110 identical protein
subunits (Galvez & Castano, 1976; Goodman etal.,1977;Qazi
et. al.,2007).
3. Taxonomy-
Mungbean yellow mosaic virus, MungbeanyellowmosaicIndia
virus, Horsegram yellow mosaic virus and Dolichos
yellow mosaic virus are bipartite begomovirus that are
responsible for causing yellow mosaic diseases in legumes
across southern Asia. MYMIV is a bipartite isolate of
Begomovirus that is more widespread in tropical and
subtropical climates (Chakraborty et al., 2003; Usharani et
al., 2005; John et al., 2008;Fazeli et al., 2009; Haq et al.,
2011a, 2011b).
4. Genome-
Genome of begomovirus is bipartite (Honda and Ikegami
1986; Vanitharani et al.1996; Mandal et al. 1997;
Karthikeyan et al. 2004).Both the virions and
complementary sense standards have coding regions,
positive and negative respectively. Double standard
intermediates are responsible for replication of the genome
by rolling circle replication wherecomponentBisdependent
on component A for replication with a common region
consisting of two divergent promoters differentially
regulating temporal expression of viral genes (SIB 2008).
5. Population genomics-
Experiments on global scale population structure of
begomovirus concluded seven major sub populations that
can be further divided into 34 entirely different yet
genetically close minor sub populations(Prasanna et.
al.,2010).
6. Gene expression-
Common region is responsible forbidirectionaltranscription
in begomovirus while protein expression takes place at
subgenomic RNA level. Bipartite begomovirus majorly a
disease causing mesobiotic pathogen in legumes has two
components- Component A encodes six proteins CP(on v-
sense) & Rep, TrAP/AL2, REn, AC4, AV2(on c-sense) and
component B encodes2 proteinsBV1(on v-sense) & BC1(on
c-sense) both involved in movement (Sunter and Bisaro
1992, Bisaro 2006, Sunter and Bisaro 1992, Noueiry et al.
International Research Journal of Engineering and Technology (IRJET) e-ISSN: 2395-0056
Volume: 05 Issue: 08 | Aug 2018 www.irjet.net p-ISSN: 2395-0072
Β© 2018, IRJET | Impact Factor value: 7.211 | ISO 9001:2008 Certified Journal | Page 1866
1994, Fontes et al.1994, Laufs et al. 1995, Sanderfoot and
Lazarowitz 1996, Hanley-Bowdoin et al. 2000, Arguello-
Astorga et al. 2004, Bisaro 2006, SIB 2008, Hanley-Bowdoin
et al. 2013, ICTV 2017).
7. Economic Impact-
After first report of yellow mosaic disease on major pulse
group,its economic impact was expressed by various
scientists.Yellow mosaic disease has a potential to cause 85-
100% yield loss in black gram and mungbean
(Nene,1973).Yellow mosaic disease accountedforatotalloss
of 105,000 metric tonnes in soybean alone(Wrather et.
al.,1997).Total yield losses in blackgram,soybean and
mungbean collectively are expected to be around $ 300
million per year(Verma and Malathi,2003).
8. Host Range-
Begomovirus are reported to hold a wider host range but in
general it serves on dicotyledonous plant.Specific major
legume genera includes Phaseolus, Vigna, Macroptilium,
Calopogonium(Bird et al., 1972; Bird,Sanchez&Vakili,1973;
Meiners et al., 1973).Rarely infected legume genera are
Cassia, Cajanus, Glycine, Rhynchosia and Phaseolus (J.
Bird).Intrinsic research work led to identify Limabean or
lablab bean(Capoor and Verma,1948),Mungbean,urdbean
and cowpea (Nariyani,1960;Nene,1973), soybean
(Suteri,1974,Fernandeset al. 2009),horsegram (Muniyappa
et al., 1975),Frenchbean (Singh, 1979), wild bush bean or
quail bean (Macroptillium lathyroides) (Lima et al. 2013).
9. Transmission-
Original transmission was reported by whitefly Bemisia
tabaci (Hemiptera: Aleyrodidae) in a persistent, non-
propagative manner across the globe(Costa,1965). Bemisia
tabaci race sidae is known to spread the virus(Bird et.
al.,1972).Both males and females are capable to act as a
vector of the virus and no evidence of larvae acting as
vectors has been found(Bird et. al.,1973).Infectionispossible
in the hosts within six days of acquisition and
inoculation(Bird,1973). However longer feedingperiodsare
needed to spread the disease effectively.16-21 days
approximately is the period to act as potential vectors after
acquisition period of 10-15 minutes (Gamez, 1971; Bird et
al., 1973). All whiteflies sub types B. tabaci MEAM1 (Middle
East-Asia Minor 1) (Biotype B), B. tabaci NW (New World)
(Biotype A), B. tabaci NW2 (New World 2), and B. tabaci
MED (Mediterranean; Biotype Q) (Barbosa et al. 2015) are
responsible for transmission of virusexceptbiotype-bwhich
acts as a vector only on monocots.Circulative transmissionis
known not to occur but replication of the virus inside the
vector is questionable (Rosen et. al.,2015).
10. Molecular characterization-
Molecular characterization and infectivity test led to the
conclusion that viral isolates obtained from Indian cowpea
were 94-98% similar in its genomic sequence of DNA-A and
DNA-B and shared a close relationship with the other
isolates obtained so far, thus proving the Koch’s postulates
for begomovirus association with mungbean yellow mosaic
disease-India in mungbean and cowpea(Singh et. al.,2011).
11. Antigenicity
All the begomoviruses were put to test and serological tests
evaluated them to be closely related. HarrisonandRobinson,
1999 used monoclonal antibodies to group various
begomoviruses geographically on the basis of shared
epitopes.
12. Infectivity-
YMD’s first incidence was marked in western and northern
India in lima bean(Capoor and Verma,1948). Nariyani,1960
wasthe first to report yellow mosaic disease in mungbeanin
Indain subcontinent. Infection in legumesarecausedbybean
golden mosaic viruswhich is a type strain of begomovirus.It
includes Mungbean yellow mosaic India virus, Mungbean
yellow mosaic virus, Dolichos yellow mosaic virus which
have bipartite genome and produce mosaic and leaf
yellowing as a the major symptoms in legumes(Varma and
Malathi 2003 ; Balaji et al. 2004; Girish et al.
2005).Furthermore, Mungbean yellow mosaic India virus
(MYMIV) and Mungbean yellow mosaic virus (MYMV) were
reported to be more virulent,having more hosts and
abundant(Fauquet and Stanley,2003) and the other two
strains Dolichos yellow mosaic virus and Horsegram yellow
mosaic virus are specific and rare(Maruthi et. al.,2005).
Molecular characterization proved begomovirus subgroup-
III isolates were responsible to cause mild mosaic infection
on Vigna mungo var. Sylvestris L.(Naimuddin et. al.,2011).
13. Management-
For the management of begomovirus it is important to lower
down the inoculum and and the vector population.
Cutting and destruction of infected portion of the plant in
case of low or moderate infection and uprootingof the plant
in case of severe infection is profitable.
Seed treatment with imidachlorpid @ 5ml/kg seed withtwo
sprays of imidachlorpid @ 0.5 ml/l at 25 and 40 DAS were
found to be effective against the infection (Jayappa et.
al.,2017). Another effective method is two sprays of
neemazal @ 3ml/l after seed treatment with imidachlorpid
@ 5ml/kg seed (Jayappa et. al.,2017). Predators offer a
better possibility of vector control. These predators include
lacewings, bigeyed bugs, and minute pirate bugs. Lady
beetlesincluding Clitostethus arcuatus (on ash whitefly),the
Asian multicolored lady beetle Harmonia axyridis and scale
predators, such as Scymnus or Chilocorus species feed on
whiteflies (UCIPM).
International Research Journal of Engineering and Technology (IRJET) e-ISSN: 2395-0056
Volume: 05 Issue: 08 | Aug 2018 www.irjet.net p-ISSN: 2395-0072
Β© 2018, IRJET | Impact Factor value: 7.211 | ISO 9001:2008 Certified Journal | Page 1867
14. Conclusion and Future Prospects-
Begomovirus with their quantity,infectivity and strain
differentiations have always attaracted us to conduct
exemplary research on them. The wide host range they
share, the virulence they have developed and their close
association with vector whitefly has brought them to
immediate research action and work at present. Also as it
fears nutritional security in vegetables like okra, brinjal,
tomato, chili, cassava and food security throughlegumesand
cereals, it has come up as an important and immediate
research interest. Disease symptoms and particle
morphology suggest that bean golden mosaic
viruses(subgroup-III of begomovirus) from Puerto Rico, El
Salvador, Colombia, Guatamala, and probably Brazil are the
same virus, although possible strain relationships are not
worked out. Causal agents of similar diseases in other
tropical areas have not been characterized. Thus, future
research should be emphasized to workout strain
relationships and solve complexity of gene order of
begomovirus. Another challenge is to markthecommongene
pool,if any,of begomovirus to understand population
distribution and gene flow to check increasing levels of
virulent strains. The disease causing ability of the
begomovirus has increased due to evolutionofmorevirulent
strains and is likely to increase due to uncontrollable spread
of whitefly population, tropical climate and widespread
cultivation of legumes. Hence, Integrated approach towards
the solution must be adopted to control the same. B. tabaci
MEAM1 (Middle East-Asia Minor 1) (Biotype B) is found to
violate rule boundaries of begomovirus and is found to
transmit virus on monocots like maize which can later
spread to legumes owing new threat to food security of
future. Thus, thrust should be laid to limit strain of
begomovirus on monocots. Extreme research work is also
demanded in the field of protein mapping technology to
mark both protein in a bipartite virus and understand their
structure to break gene bond which would eventually
control the disease spread. A new approach .i.e. application
of new generation sequencing (NGS) technology for better
diagnosis by metagenomic analysis and deep sequencing is
yet to be carried out in begomovirus. This will allow us to
develop better understanding of viruses and their
associations and control them to achieve food security.
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IRJET- Research Developments in Begomovirus in Legumes : Past Achievements, Present Scenario and Future Thrust Areas

  • 1. Β© 2018, IRJET | Impact Factor value: 7.211 | ISO 9001:2008 Certified Journal | Page 1865 Research developments in begomovirus in legumes : Past chievements, Present scenario and Future thrust areas Lavania Pranesh1, Singh Priya2, Sharma Manju3 1&2Department of Plant Pathology, College of Agriculture, G.B. Pant University of Agriculture and Technology, Pantnagar, Uttarakhand, India 3Assistant Professor, Department of Plant Pathology, College of Agriculture, G.B. Pant University of Agriculture and Technology, Pantnagar, Uttarakhand, India ---------------------------------------------------------------------***--------------------------------------------------------------------- Abstract - Begomovirus – A well recognized genus of plant viruses classified under Geminiviridae family of Group II (single stranded circular DNA) is known to cause virulence in a large range of hosts including tomato, okra, cotton, cassava, bittergourd, chilli, croton, cucumber, eggplant, jatropha, mentha, mesta, papaya, potato, tobacco, legumes and many more. It is believed that organism has caused major economic losses on cropsof all types. However,owing to today’s demand of increased production, land area and productivity of pulses for future food and nutritional security, this paper will summarize collectively the research study and experiments conducted on legumes till date, present research work being conducted and future thrust areas on begomoviruses w.r.t. legumes like common bean (Phaseolus vulgaris), mung bean (Vigna radiata), urd bean (Vigna mungo), pigeon pea (Cajanuscajan),mothbean(Vigna aconitifolia), cowpea (Vignaungiculata),velvetbean(Mucuna pruriens), frenchbean (Phaseolus vulgaris), soybean (Glysine max), long yard bean (Vigna sesquipedalis) etc. Prime focus areas of this review will include molecular characterization, DNA replication, phylogenetic analysis and infectivity patterns on host legumes. Key Words: Begomovirus, legumes, Bemisia tabaci, yellow mosaic disease, bean common mosaic virus 1. Introduction- Among all the diseases of plants, viral diseases are the most complex and the least known. Intensive research in the recent past has led to showcase some facts with clear understanding. This review will deal with morphology, taxonomy, genome, population genomics, gene expression, economic impact, host range, transmission, molecular characterization, antigenicity, infectivityandmanagementof begomoviruses that forms largest group of disease causing viruses of plants and specifically of legumes. 2. Morphology- Virions of geminivirus sub group III are germinate .i.e. twinned with two incomplete icosahedra. They are non- enveloped in nature and have a common dimensionof30-38 nm * 18-28 nm (length*diameter) with 22 pentameric capsomers per nucleocaspid and 110 identical protein subunits (Galvez & Castano, 1976; Goodman etal.,1977;Qazi et. al.,2007). 3. Taxonomy- Mungbean yellow mosaic virus, MungbeanyellowmosaicIndia virus, Horsegram yellow mosaic virus and Dolichos yellow mosaic virus are bipartite begomovirus that are responsible for causing yellow mosaic diseases in legumes across southern Asia. MYMIV is a bipartite isolate of Begomovirus that is more widespread in tropical and subtropical climates (Chakraborty et al., 2003; Usharani et al., 2005; John et al., 2008;Fazeli et al., 2009; Haq et al., 2011a, 2011b). 4. Genome- Genome of begomovirus is bipartite (Honda and Ikegami 1986; Vanitharani et al.1996; Mandal et al. 1997; Karthikeyan et al. 2004).Both the virions and complementary sense standards have coding regions, positive and negative respectively. Double standard intermediates are responsible for replication of the genome by rolling circle replication wherecomponentBisdependent on component A for replication with a common region consisting of two divergent promoters differentially regulating temporal expression of viral genes (SIB 2008). 5. Population genomics- Experiments on global scale population structure of begomovirus concluded seven major sub populations that can be further divided into 34 entirely different yet genetically close minor sub populations(Prasanna et. al.,2010). 6. Gene expression- Common region is responsible forbidirectionaltranscription in begomovirus while protein expression takes place at subgenomic RNA level. Bipartite begomovirus majorly a disease causing mesobiotic pathogen in legumes has two components- Component A encodes six proteins CP(on v- sense) & Rep, TrAP/AL2, REn, AC4, AV2(on c-sense) and component B encodes2 proteinsBV1(on v-sense) & BC1(on c-sense) both involved in movement (Sunter and Bisaro 1992, Bisaro 2006, Sunter and Bisaro 1992, Noueiry et al. International Research Journal of Engineering and Technology (IRJET) e-ISSN: 2395-0056 Volume: 05 Issue: 08 | Aug 2018 www.irjet.net p-ISSN: 2395-0072
  • 2. Β© 2018, IRJET | Impact Factor value: 7.211 | ISO 9001:2008 Certified Journal | Page 1866 1994, Fontes et al.1994, Laufs et al. 1995, Sanderfoot and Lazarowitz 1996, Hanley-Bowdoin et al. 2000, Arguello- Astorga et al. 2004, Bisaro 2006, SIB 2008, Hanley-Bowdoin et al. 2013, ICTV 2017). 7. Economic Impact- After first report of yellow mosaic disease on major pulse group,its economic impact was expressed by various scientists.Yellow mosaic disease has a potential to cause 85- 100% yield loss in black gram and mungbean (Nene,1973).Yellow mosaic disease accountedforatotalloss of 105,000 metric tonnes in soybean alone(Wrather et. al.,1997).Total yield losses in blackgram,soybean and mungbean collectively are expected to be around $ 300 million per year(Verma and Malathi,2003). 8. Host Range- Begomovirus are reported to hold a wider host range but in general it serves on dicotyledonous plant.Specific major legume genera includes Phaseolus, Vigna, Macroptilium, Calopogonium(Bird et al., 1972; Bird,Sanchez&Vakili,1973; Meiners et al., 1973).Rarely infected legume genera are Cassia, Cajanus, Glycine, Rhynchosia and Phaseolus (J. Bird).Intrinsic research work led to identify Limabean or lablab bean(Capoor and Verma,1948),Mungbean,urdbean and cowpea (Nariyani,1960;Nene,1973), soybean (Suteri,1974,Fernandeset al. 2009),horsegram (Muniyappa et al., 1975),Frenchbean (Singh, 1979), wild bush bean or quail bean (Macroptillium lathyroides) (Lima et al. 2013). 9. Transmission- Original transmission was reported by whitefly Bemisia tabaci (Hemiptera: Aleyrodidae) in a persistent, non- propagative manner across the globe(Costa,1965). Bemisia tabaci race sidae is known to spread the virus(Bird et. al.,1972).Both males and females are capable to act as a vector of the virus and no evidence of larvae acting as vectors has been found(Bird et. al.,1973).Infectionispossible in the hosts within six days of acquisition and inoculation(Bird,1973). However longer feedingperiodsare needed to spread the disease effectively.16-21 days approximately is the period to act as potential vectors after acquisition period of 10-15 minutes (Gamez, 1971; Bird et al., 1973). All whiteflies sub types B. tabaci MEAM1 (Middle East-Asia Minor 1) (Biotype B), B. tabaci NW (New World) (Biotype A), B. tabaci NW2 (New World 2), and B. tabaci MED (Mediterranean; Biotype Q) (Barbosa et al. 2015) are responsible for transmission of virusexceptbiotype-bwhich acts as a vector only on monocots.Circulative transmissionis known not to occur but replication of the virus inside the vector is questionable (Rosen et. al.,2015). 10. Molecular characterization- Molecular characterization and infectivity test led to the conclusion that viral isolates obtained from Indian cowpea were 94-98% similar in its genomic sequence of DNA-A and DNA-B and shared a close relationship with the other isolates obtained so far, thus proving the Koch’s postulates for begomovirus association with mungbean yellow mosaic disease-India in mungbean and cowpea(Singh et. al.,2011). 11. Antigenicity All the begomoviruses were put to test and serological tests evaluated them to be closely related. HarrisonandRobinson, 1999 used monoclonal antibodies to group various begomoviruses geographically on the basis of shared epitopes. 12. Infectivity- YMD’s first incidence was marked in western and northern India in lima bean(Capoor and Verma,1948). Nariyani,1960 wasthe first to report yellow mosaic disease in mungbeanin Indain subcontinent. Infection in legumesarecausedbybean golden mosaic viruswhich is a type strain of begomovirus.It includes Mungbean yellow mosaic India virus, Mungbean yellow mosaic virus, Dolichos yellow mosaic virus which have bipartite genome and produce mosaic and leaf yellowing as a the major symptoms in legumes(Varma and Malathi 2003 ; Balaji et al. 2004; Girish et al. 2005).Furthermore, Mungbean yellow mosaic India virus (MYMIV) and Mungbean yellow mosaic virus (MYMV) were reported to be more virulent,having more hosts and abundant(Fauquet and Stanley,2003) and the other two strains Dolichos yellow mosaic virus and Horsegram yellow mosaic virus are specific and rare(Maruthi et. al.,2005). Molecular characterization proved begomovirus subgroup- III isolates were responsible to cause mild mosaic infection on Vigna mungo var. Sylvestris L.(Naimuddin et. al.,2011). 13. Management- For the management of begomovirus it is important to lower down the inoculum and and the vector population. Cutting and destruction of infected portion of the plant in case of low or moderate infection and uprootingof the plant in case of severe infection is profitable. Seed treatment with imidachlorpid @ 5ml/kg seed withtwo sprays of imidachlorpid @ 0.5 ml/l at 25 and 40 DAS were found to be effective against the infection (Jayappa et. al.,2017). Another effective method is two sprays of neemazal @ 3ml/l after seed treatment with imidachlorpid @ 5ml/kg seed (Jayappa et. al.,2017). Predators offer a better possibility of vector control. These predators include lacewings, bigeyed bugs, and minute pirate bugs. Lady beetlesincluding Clitostethus arcuatus (on ash whitefly),the Asian multicolored lady beetle Harmonia axyridis and scale predators, such as Scymnus or Chilocorus species feed on whiteflies (UCIPM). International Research Journal of Engineering and Technology (IRJET) e-ISSN: 2395-0056 Volume: 05 Issue: 08 | Aug 2018 www.irjet.net p-ISSN: 2395-0072
  • 3. Β© 2018, IRJET | Impact Factor value: 7.211 | ISO 9001:2008 Certified Journal | Page 1867 14. Conclusion and Future Prospects- Begomovirus with their quantity,infectivity and strain differentiations have always attaracted us to conduct exemplary research on them. The wide host range they share, the virulence they have developed and their close association with vector whitefly has brought them to immediate research action and work at present. Also as it fears nutritional security in vegetables like okra, brinjal, tomato, chili, cassava and food security throughlegumesand cereals, it has come up as an important and immediate research interest. Disease symptoms and particle morphology suggest that bean golden mosaic viruses(subgroup-III of begomovirus) from Puerto Rico, El Salvador, Colombia, Guatamala, and probably Brazil are the same virus, although possible strain relationships are not worked out. Causal agents of similar diseases in other tropical areas have not been characterized. Thus, future research should be emphasized to workout strain relationships and solve complexity of gene order of begomovirus. Another challenge is to markthecommongene pool,if any,of begomovirus to understand population distribution and gene flow to check increasing levels of virulent strains. The disease causing ability of the begomovirus has increased due to evolutionofmorevirulent strains and is likely to increase due to uncontrollable spread of whitefly population, tropical climate and widespread cultivation of legumes. Hence, Integrated approach towards the solution must be adopted to control the same. B. tabaci MEAM1 (Middle East-Asia Minor 1) (Biotype B) is found to violate rule boundaries of begomovirus and is found to transmit virus on monocots like maize which can later spread to legumes owing new threat to food security of future. Thus, thrust should be laid to limit strain of begomovirus on monocots. Extreme research work is also demanded in the field of protein mapping technology to mark both protein in a bipartite virus and understand their structure to break gene bond which would eventually control the disease spread. A new approach .i.e. application of new generation sequencing (NGS) technology for better diagnosis by metagenomic analysis and deep sequencing is yet to be carried out in begomovirus. This will allow us to develop better understanding of viruses and their associations and control them to achieve food security. REFERENCES 1. Arguello-Astorga G, Lopez-Ochoa L, KongLJ,Orozco BM, Settlage SB, Hanley-Bowdoin L 2004 A novel motif in geminivirus replication proteins interacts with the plant retinoblastoma-related protein J Virol. 2004 May; 78(9) 4817-26. 2. 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