ORIGINAL ARTICLE
Recombination Among Begomoviruses on Malvaceous Plants
Leads to the Evolution of Okra Enation Leaf Curl Virus in Pakistan
Saad Serfraz
1
, Imran Amin
1
, Khalid P. Akhtar
2
and Shahid Mansoor
1
1 Agricultural Biotechnology Division, National Institute for Biotechnology and Genetic Engineering, Faisalabad, Pakistan
2 Plant Protection Division, Nuclear Institute for Agriculture and Biology, Faisalab ad, Pakistan
Keywords
begomoviruses, malvaceous host, new
species, Okra enation leaf curl virus
Correspondence
S. Mansoor, National Institute for
Biotechnology and Genetic Engineering,
Faisalabad, Pakistan.
E-mail: Shahidmansoor7@gmail.com
Received: August 10, 2014; accepted:
December 29, 2014.
doi: 10.1111/jph.12373
Abstract
Whitefly transmitted begomoviruses (family Geminiviridae) are the major
reason for significant yield losses of dicotyledonous crops in tropics and
subtropics. Okra (Abelmoschus esculentus) is one of the important vegetable
crops, and leaf curl disease caused by geminiviruses is the most important
limiting factor for its production in Pakistan. Here, we report a new species
of okra-infecting begomovirus in south-eastern region of Pakistan and the
name Okra enation leaf curl virus (OELCuV) complex is proposed. This okra
enation leaf curl disease complex (OELCuD) in Pakistan is found to be
associated with Ageratum conyzoides symptomless alphasatellite (ACon-
SLA). All efforts to clone the betasatellite were unsuccessful. Comprehen-
sive sequence analyses suggest that intermalvaceous recombination
between okra and cotton-infecting begomoviruses resulted in the evolu-
tion of the new species. Surprisingly, Bhendi yellow vein mosaic virus (BY-
VMV) which has not been reported previously from Pakistan is the major
parent while Cotton leaf curl Multan virus (CLCuMV) acts as a distant parent
of the virus. Comparative recombination analysis also reveals that okra-
infecting begomoviruses from south and north-western India is causing
OELCuD in the Pakistan by recombining with CLCuMV at the Rep (1964
1513 nts). Recombination is common among geminiviruses and recom-
bining of BYVMV and CLCuMV resulted in a new species: OELCuV. To
the best of our knowledge, this evolution of a new species of okra-infect-
ing begomovirus is the first report of intermalvaceous recombination
where Rep acts as the target region.
Introduction
Plant viruses belonging to the family Geminiviridae are
with single-stranded (ss) DNA genomes, encapsidated
within two incomplete icosahedral particles inter-
twined to form a unique type of morphology of viral
coat protein with a diameter ~18 nm and length
~30 nm (Fauquet et al. 2008). According to recent
classification, the family Geminivirdae is divided into
seven different genera (Mastrevirus, Topocuvirus, Curto-
virus, Becurtovirus, Turncurtovirus, Eragovirus and
Begomovirus) based on the genome organization, insect
vector and host range (Adams et al. 2013). Begomovi-
rus is the largest and economically the most important
genus within the family. Begomoviruses have a wide
host range which includes economically important
crops from diverse families such as Malvaceae, Solana-
ceae, Rosaseaece and Fabaceae and is transmitted by
whitefly Bemisia tabaci (Gennadius; Varma and Mala-
thi 2003; Akhtar et al. 2014; Ullah et al. 2014, 2015).
Begomoviruses genome organization differs according
to geographical location. The genome of viruses native
to the New World (NW) comprises two components,
DNA-A and DNA-B. With few exceptions, most of the
Old World (OW) begomoviruses are monopartite and
show a synergistic association with a helper compo-
nent (betasatellite, alphasatellites; Mansoor et al.
2000; Briddon et al. 2004; Briddon and Stanley 2006).
J Phytopathol 163 (2015) 764–776 Ó 2015 Blackwell Verlag GmbH764
J Phytopathol
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Supporting Information
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Figures
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Figure 1 | Phylogeographic analysis of OELCuV-Pk (HG518793, HG 938358, HG 938359, HG938362). Colour distribution shows geographical location of each isolate.
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Figure 2 | Neighbour-joining-based phylogenetic tree based on the alignment of alphasatellites of OELCuD Neo-23 to Neo-27 (HG518788-HG518792) with different species alphasatellites available in database. Numbers on node show the bootstrap confidence values (1000 replicates). Tree is rooted with Cotton leaf curl Gezira betasatellite (CLCuGB). Sequences include Cotton leaf curl Burewala alphasatellite (CLCuBuA), Cotton leaf Curl Multan alphasatellite (CLCuMA), Papaya leaf Curl alphasatellite (PaLCuA) and Ageratum conyzoides associated symptomless alphasatellite (AConSLA).
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Figure 3 | RDP analysis showing recombination between BYVMV and OELCuV. Part a shows the recombination event with its major (BYVMV) and minor (OELCuV-IN) parent. Part b shows the graphical representation of recombination event. Table at the bottom shows the values of algorithms used by RDP 4.
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Figure 4 | Plotcon analysis, (a) shows the graph between OELCuV-PK and OELCuV-IN. (GU11997). (b) Shows the graph between OELCuV and BYVMV (JQ35953).
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Figure 5 | Phylogenetic dendrogram showing comparison of recombinant fragment from 1964 to 1513 nts of Rep protein with related sequences present in the database. OELCuV-PK, BYVMV.
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Figure 6 | RDP analysis showing recombination between MYVMV (FJ345400) and CLCuMV (AJ496287) producing an entire REP.
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