ISSN 1022-7954, Russian Journal of Genetics, 2007, Vol. 43, No. 4, pp. 381–386. © Pleiades Publishing, Inc., 2007.
Original Russian Text © T.V. Danilova, M.Yu. Kuklev, G.N. Andreeva, V.S. Shevelukha, G.I. Karlov, 2007, published in Genetika, 2007, Vol. 43, No. 4, pp. 482–488.
381
INTRODUCTION
Wild silverleaf sunflower, Helianthus agrophyllus,
is resistant to a number of diseases and parasites that
affect cultivated sunflower, and is used in breeding pro-
grams as a source of the resistance to Phomopsis stem
canker, Diaporthe helianthi [1, 2], white mold, Sclero-
tinia sclerotiorum [3], downy mildew, Plasmopara hal-
stedii [4], sunflower broomrape, Orobanche cumana
[5], and drought [6]. However, breeding of resistant
forms of sunflower is restrained by the lack of effective
methods of selection for resistance. In recent years,
molecular marking of economically important traits of
cultivated plants, including the resistance to different
pathogens, has widely used.
At present, resistance genes have been studied in
many plants [7]. The resistance gene products (R-pro-
teins) of monocotyledonous and dicotyledonous plants
have conservative domains, the structures of which are
similar, albeit the differences in the mechanisms of
infection with pathogens and parasites. Depending on
the presence of certain conservative regions, different
classes of resistance genes are recognized. The proteins
containing N-terminal nucleotide binding site (NBS)
and C-terminal leucine-rich repeats (LRR), represent
the products of the largest in number class of the resis-
tance genes studied so far (NBS–LRR). The NBS–LRR
proteins in turn are subdivided into subclasses in accor-
dance with the N terminus structure. One of the sub-
classes includes the proteins with TIR domains, homol-
ogous to Toll receptor protein of Drosophila and human
interleukin-1 receptor (TIR NBS–LRR). These proteins
were found only in dicotyledons. They include the
products of the N gene of tobacco, L6 gene of flax, and
RPP5 gene of Arabidopsis. The second subclass com-
prises the proteins without TIR domain (NBS–LRR).
These proteins were described both in monocotyledons
and dicotyledons, including, for example, the products
of the Rx gene of potato and the Mlo gene of barley.
Proteins of the third subclass are characterized by the
presence of a leucine zipper structure in N-terminal
region (LZ) (LZ–NBS–LRR). These are the products of
the RPS2 gene of Arabidopsis, and Mi and Prf genes of
tomato [8, 9].
Based on conserved amino acid sequences of NBS
and LRR domains, degenerate primers were designed.
PCR with these primers produced DNA sequences of
putative resistance genes in many plant species. These
sequences are called resistance-gene analogs (RGA) or
resistance candidate genes. Genetic analysis of RGA
showed that these sequences were closely linked to the
genes conferring resistance to different pathogens and
parasites. Amplified sequences can serve for the pro-
duction of molecular markers for selection of resistant
forms, or for molecular cloning of the novel resistance
gene. The method discussed provided marking and
mapping of the resistance genes in such cultivated
plants as sunflower (downy mildew) [4], potato and
tomato (Phytophtora and nematode) [10, 11], citrus
plants, Citrus and Poncirus (virus and nematode of cit-
rus) [12], melon (fusariose and aphid) [13], and soybean
(virus, Phytophtora, and powdery mildew) [14, 15].
In the present study, a possibility of using the com-
bination of primers designed from the conservative
NBS domains of the genes RPS2 of Arabidopsis, N of
tobacco, and L6 of flax, for amplification of the resis-
tance-gene analogs from silverleaf sunflower (Helian-
thus agrophyllus) was evaluated. The produced PCR
products were cloned and characterized.
Cloning and Analysis of the Resistance Gene Fragments
from Silverleaf Sunflower Helianthus agrophyllus
T. V. Danilova, M. Yu. Kuklev, G. N. Andreeva, V. S. Shevelukha, and G. I. Karlov
Timiryazev Moscow Agricultural Academy, Moscow, 127550 Russia;
fax: (495)977-84-28; e-mail: karlov@timacad.ru
Received October 10, 2005; in final form, May 5, 2006
Abstract—Using a combination of degenerate primers designed from the NBS domains of the resistance
genes, amplification and subsequent cloning of the resistance gene fragments from sunflower (Helianthus agro-
phyllus) was conducted. Sequences of cloned PCR products differed from one another and displayed homology
to NBS domain fragments of the already known plant resistance genes, as well as to the analogous genes from
different classes. The highest homology was shown to the NBS domain regions of cultivated sunflower and the
other members of the family Compositae. Two cloned fragments had open reading frames, while the other
sequences carried stop codons and seemed to belong to pseudogenes. Amino acid sequences of Helianthus
agrophyllus analyzed contained conservative regions typical of NBS domains of the resistance gene products.
DOI: 10.1134/S1022795407040059
PLANT
GENETICS