Emil Khavkin - Academia.edu (original) (raw)
Papers by Emil Khavkin
Recently new aggressive lineages have emerged in the West European populations of <em>Phyto... more Recently new aggressive lineages have emerged in the West European populations of <em>Phytophthora infestans</em> and have overcome many elite potato varieties known for high late blight resistance. These lineages have rapidly moved eastward displacing the previously known strains of the pathogen. To evaluate the changes in <em>Ph. infestans</em> populations which spread across the North-Western Russia, we hawe realized in 2013-2014 two pilot experiments with isolates obtained from leaves collected in the field potato plots of the Pushkin laboratories of the N.I. Vavilov Institute of Plant Genetic Resources (VIR) in the Leningrad Region of Russia. In this study the phytopathologic characteristics of Pushkin isolates of <em>Ph. infestans</em> were supplemented, for the first time, with the molecular evidence. Potato leaves naturally infected with <em>Ph. infestans</em> were collected from variety Sarpo Mira, which manifested high level ...
Russian Agricultural Sciences, 2014
Isolates of P. infestans were collected in 2013 from infected potato leaves in the Pushkin field ... more Isolates of P. infestans were collected in 2013 from infected potato leaves in the Pushkin field plots of the Institute of Plant Industry, St. Petersburg. The phytopathological data on A1/A2 mating types matched the evidence obtained by CAPS marker analysis. When genotyped with SSR markers, the Pushkin isolates were clearly distinct from West European strains described by Cooke et al. (2012).
Russian Agricultural Sciences, 2010
To investigate whether the presence of R genes is associated with higher late blight resistance o... more To investigate whether the presence of R genes is associated with higher late blight resistance of potato cultivars, we modified an earlier described SCAR marker for the R1 gene and developed a new SCAR marker for the R3 gene. We demonstrated the significant input of R1 transferred from Solanum demissum and probably S. stoloniferum into foliage and tuber resistance; the effect of R3 introgression was less conclusive.
Russian Journal of Plant Physiology, 2006
... Chandler, J., Corbesier, L., Spielmann, P., Dettendorfer, J., Stahl, D., Apel, K., and Melzer... more ... Chandler, J., Corbesier, L., Spielmann, P., Dettendorfer, J., Stahl, D., Apel, K., and Melzer, S., Modulating Flower-ing Time and Prevention of Pod Shatter in Oilseed Rape, Mol. ... Calonje, M., Cubas, P., Martinez-Zapater, JM, and Car-mona, MJ, Floral Meristem Identity Genes Are ...
Russian Journal of Plant Physiology, 2006
The method of polymerase chain reaction was used to amplify a fragment of the LZ-NBS-LRR receptor... more The method of polymerase chain reaction was used to amplify a fragment of the LZ-NBS-LRR receptor kinase gene R1; the gene was transferred into potato (Solanum tuberosum) from its wild-growing relative S. demissum and confers the race-specific recognition of the pathogen Phytophthora infestans. To verify this method as a test for the presence of the late blight resistance gene R1, the amplified genome fragment was cloned from the potato hybrid comprising the germplasm of S. demissum. The primary structure of this fragment, which corresponded to the receptor domain of kinase, did not practically differ from the matching sequence in S. demissum. In addition, the method was verified by scoring the set of plant differentials, wherein the presence of R1 was established with race-specific Phytophthora isolates. By screening 70 potato cultivars, we established a significant relationship between the presence of the gene R1 fragment and the phenotypic characters of late blight resistance and late maturity. This evidence supports the idea that R1 was introgressed from short-day S. demissum into potato plants together with some gene(s) conferring late transition to flowering.
Planta, 1977
The sequential appearance of a specific group of embryonal antigens (EA), presumably globulins, w... more The sequential appearance of a specific group of embryonal antigens (EA), presumably globulins, was demonstrated in developing maize (Zea mays L.) caryopses using a double immunodiffusion test with absorption of common antigens. Cross immunoelectrophoresis was employed to follow the differential pattern of EA accumulation in the growing scutellum and embryonic axis. The transient nature of two predominant EA seems to indicate their role as specific protein reserves of embryonal tissues. Another presumably organ-specific EA was maintained in callus obtained from a 28-day-old culture of scutellum isolated from the mature non-germinated caryopsis.
Biochemie und Physiologie der Pflanzen, 1982
ABSTRACT Erythrocyte-binding proteins (lectins) were found in the apical meristem, the stele and ... more ABSTRACT Erythrocyte-binding proteins (lectins) were found in the apical meristem, the stele and the cortex with epidermis of the primary root of maize seedlings, and some of these lectins were present only in the tissues possessing the vascular elements. We presumed that these vascular-specific lectins could be related to the stele-specific antigens reported previously (Khavkin et al. 1980). In the electrointmnnodiffusion test with polyvalent antisera against the stele or the cortex proteins, proteins solubilized from human erythrocytes preagglutinated with the stele extract produced two common antigens and three stele-specific precipitin bands. To investigate these lectins in more detail, the antiserum was raised in rabbits against stelar proteins immobilized on the rabbit erythrocyte ghosts. This antiserum precipitated up to five antigens in one-dimensional immunoelectropherograms of the root extracts, including the vascular-specific lectins absent from the cortex. One of the stele-specific ectins was apparently a precipitin-like lectin interacting with non-immune serum proteins.
Russian Journal of Plant Physiology, 2000
Russian Journal of Plant Physiology, 2005
The arabidopsis gene LEAFY controls the induction of flowering and maintenance of the floral meri... more The arabidopsis gene LEAFY controls the induction of flowering and maintenance of the floral meristem identity. By comparing the primary structure of LEAFY and its homologs in other Brassicaceae species and beyond this family, we singled out four clusters corresponding to three systematically remote families of angiosperms, Brassicaceae, Solanaceae, and Poaceae, and to gymnosperms. Both structural and functional distinctions of LEAFY homologs from their arabidopsis prototype expanded in the range Brassicaceae-Solanaceae-Poaceae. A LEAFY homolog from B. juncea cloned in our laboratory was used as a hybridization probe to analyze the restriction fragment length polymorphism in six Brassica species comprising diploid (AA, BB, and CC) and allotetraploid (AABB, AACC, and BBCC) genomes. In this way we recognized LEAFY fragments specific of genomes A, B, and C; in contrast, the variations of the length and structure of the LEAFY intron 2 were not genome-specific. LEAFY polymorphism in the Brassica accessions comprising genome B was related to their geographic origin and apparently to the adaptation to day length.
Zeitschrift für Pflanzenphysiologie, 1978
Antigens and isozyme spectra characteristic of epigenotypes of specialized cells in intact plants... more Antigens and isozyme spectra characteristic of epigenotypes of specialized cells in intact plants were preserved in successive subcultures of the four types of callus tissues derived from mesocotyls and scutella of inbreds CI03 and W155 and their crosses.
Zeitschrift für Pflanzenphysiologie, 1979
A shake flask culture was established from maize (hybrid C 103 X W 155) callus initiated at nodal... more A shake flask culture was established from maize (hybrid C 103 X W 155) callus initiated at nodal and internodal stem segments and maintained in the medium containing the inorganic constituents of MURASHIGE and SKOOG (1962), 20 gil sucrose, 0.4 mgll thiamine, 1 gil casein hydrolysate and 1 mgll 2,4-D. Small aggregates that lacked inner organization were predominant in the suspension. Cells were of a parenchyma-type, mostly round-shaped, and their sizes were distributed in log-normal pattern (the mean diameter 40-50 ft). The exponential growth in the batch-grown culture lasted to 4th-5th day, the transition to the stationary phase took place during the 5th-6th day, and after the 9th day a considerable degradation was observed in the culture. Doubling periods for the cell number, and dry matter and protein during the exponential growth were about 28 h at 26 DC, and the total increase was at least IS-fold during the 7-day subculture cycle. Respiration rate on the fresh weight, protein and cell basis was constant from the 1st to the 6th day and then declined 2-fold, however, it was completely restored during the first 12 h after cell transfer to the fresh medium.
Planta, 1980
A group of antigenically distinct proteins characteristic for the tissue complex of the vascular ... more A group of antigenically distinct proteins characteristic for the tissue complex of the vascular cylinders was found in maize (Zea mays L.) seedlings using an immunofiltration technique. Specific stelar antigens present in the fully developed stele (vascular cylinder) of the primary root were also found in steles extracted from adventitious roots and from the mesocotyl but were absent, within the limits of sensitivity of the immunodiffusion tests employed, in root cortex and epidermis. Some of the stelar antigens were also evident in the meristem of the primary root and were present in traces in the scutellum, the mesocotyl node, and the primary leaves plus coleoptile. The specific stelar antigens could be traced in 13-and 15-day-old developing embryos and were definitely expressed by the 21st day after pollination. Several stelar-specific antigens were found in embryo-derived callus tissues and in stem-derived cells maintained in serial suspension culture. Higher resolution of the stelar antigens by a modified technique of crossed immunoelectrophoresis was used to demonstrate several minor stelar antigens that were presumably characteristic exclusively of the completely differentiated stele. This technique along with sequential immunoprecipitation of labelled proteins provided a semiquantitative estimate of the specific stelar antigens in the meristem and the stele of the primary root, and in suspension-cultured cells which were devoid of noticeable signs of vascular differentiation.
Russian Journal of Plant Physiology, 1999
Solanum bulbocastanum comprising a CC-NBS-LRR gene RB/Rpi-blb1 confers broad-spectrum resistance ... more Solanum bulbocastanum comprising a CC-NBS-LRR gene RB/Rpi-blb1 confers broad-spectrum resistance to Phytophthora infestans and is currently employed in potato breeding for durable late blight (LB) resistance. Genomes of several Solanum species were reported to contain RB homologues with confirmed broad-spectrum defence function. With the discovery that novel P. infestans races break LB resistance mediated by the genes of broad-spectrum specificity, pyramiding several RBlike genes from various Solanum species in a single potato cultivar seems a promising approach to durable LB resistance. Here we report early evidence on RB-like sequences in the wide range of Solanum species section Petota. The panel of Solanum species was screened with three RBrelated PCR markers. RB-like sequences were found in every tested Solanum accession suggesting universal distribution of RB structural homologues among Solanum genomes, while the marker RB-629 corresponding to the RB gene was found in 14 speci...
SUMMARY Allelic diversity of the IpiO gene encoding the effector recognised by Rpi‐blb1/RB gene f... more SUMMARY Allelic diversity of the IpiO gene encoding the effector recognised by Rpi‐blb1/RB gene from S. bulbocastanum and S. stoloniferum was assessed by PCR screening and sequencing in 15 races (1, 3, 4, 10, 11, 1.2, 1.3, 1.4, 2.4, 3.4, 1.11, 1.2.3, 1.2.4, 1.3.4, and 1.2.3.4) maintained in the collection of the Institute of Phytopathology. IpiO locus was present in all these isolates. Phylogenetic analysis demonstrated that IpiO alleles from races 3 and 4 belonged to the classes I and II recognised by R pi-blb 1/RB. It follows that these “simple” races exhibit simple virulence patterns on potato comprising only R1-R11 genes from S. demissum . Growing interest in exploiting exotic Solanum germplasm in potato breeding is an incentive to reconsider and unambiguously redefine the terms “simple” and “complex” race. KEYWORDS Phytophthora infestans , Solanum , potato, late blight resistance, effectors, simple races INTRODUCTION Approaches to describe the diversity of P. infestans Various ...
Sel'skokhozyaistvennaya Biologiya, Jun 1, 2016
The authors are grateful to the Center for Collective Use of Equipment «Biotechnology» (All-Russi... more The authors are grateful to the Center for Collective Use of Equipment «Biotechnology» (All-Russian Research Institute of Agricultural Biotechnology) for sequencing Avr clones and separating SSR amplicons by capillary electrophoresis. Supported by the grants from Russian Foundation for Basic Research (projects 14-04-31613а and 16-04-00098) and the Ministry of Education and Science of the Russian Federation (project RFMEFI62114X0003). The phytopathological evaluation of Phytophthora infestans isolates was performed as part of the State Commission 0598-2015-0018.
Breeding potato for durable resistance to late blight (LB) greatly benefits from expanding the re... more Breeding potato for durable resistance to late blight (LB) greatly benefits from expanding the resource of resistance genes provided by wild Solanum species. SCAR markers for race-specific CC-NBS-LRR resistance genes (R genes) have been employed to screen Solanum accessions representing six series of section Petota. Structural homologues of particular genes were found in many taxonomically distant species and widely differed in their distribution patterns. Such evidence suggests that the R-gene structures evolved before the divergence of genomes A and B and preceded Solanum speciation.
Recently new aggressive lineages have emerged in the West European populations of <em>Phyto... more Recently new aggressive lineages have emerged in the West European populations of <em>Phytophthora infestans</em> and have overcome many elite potato varieties known for high late blight resistance. These lineages have rapidly moved eastward displacing the previously known strains of the pathogen. To evaluate the changes in <em>Ph. infestans</em> populations which spread across the North-Western Russia, we hawe realized in 2013-2014 two pilot experiments with isolates obtained from leaves collected in the field potato plots of the Pushkin laboratories of the N.I. Vavilov Institute of Plant Genetic Resources (VIR) in the Leningrad Region of Russia. In this study the phytopathologic characteristics of Pushkin isolates of <em>Ph. infestans</em> were supplemented, for the first time, with the molecular evidence. Potato leaves naturally infected with <em>Ph. infestans</em> were collected from variety Sarpo Mira, which manifested high level ...
Russian Agricultural Sciences, 2014
Isolates of P. infestans were collected in 2013 from infected potato leaves in the Pushkin field ... more Isolates of P. infestans were collected in 2013 from infected potato leaves in the Pushkin field plots of the Institute of Plant Industry, St. Petersburg. The phytopathological data on A1/A2 mating types matched the evidence obtained by CAPS marker analysis. When genotyped with SSR markers, the Pushkin isolates were clearly distinct from West European strains described by Cooke et al. (2012).
Russian Agricultural Sciences, 2010
To investigate whether the presence of R genes is associated with higher late blight resistance o... more To investigate whether the presence of R genes is associated with higher late blight resistance of potato cultivars, we modified an earlier described SCAR marker for the R1 gene and developed a new SCAR marker for the R3 gene. We demonstrated the significant input of R1 transferred from Solanum demissum and probably S. stoloniferum into foliage and tuber resistance; the effect of R3 introgression was less conclusive.
Russian Journal of Plant Physiology, 2006
... Chandler, J., Corbesier, L., Spielmann, P., Dettendorfer, J., Stahl, D., Apel, K., and Melzer... more ... Chandler, J., Corbesier, L., Spielmann, P., Dettendorfer, J., Stahl, D., Apel, K., and Melzer, S., Modulating Flower-ing Time and Prevention of Pod Shatter in Oilseed Rape, Mol. ... Calonje, M., Cubas, P., Martinez-Zapater, JM, and Car-mona, MJ, Floral Meristem Identity Genes Are ...
Russian Journal of Plant Physiology, 2006
The method of polymerase chain reaction was used to amplify a fragment of the LZ-NBS-LRR receptor... more The method of polymerase chain reaction was used to amplify a fragment of the LZ-NBS-LRR receptor kinase gene R1; the gene was transferred into potato (Solanum tuberosum) from its wild-growing relative S. demissum and confers the race-specific recognition of the pathogen Phytophthora infestans. To verify this method as a test for the presence of the late blight resistance gene R1, the amplified genome fragment was cloned from the potato hybrid comprising the germplasm of S. demissum. The primary structure of this fragment, which corresponded to the receptor domain of kinase, did not practically differ from the matching sequence in S. demissum. In addition, the method was verified by scoring the set of plant differentials, wherein the presence of R1 was established with race-specific Phytophthora isolates. By screening 70 potato cultivars, we established a significant relationship between the presence of the gene R1 fragment and the phenotypic characters of late blight resistance and late maturity. This evidence supports the idea that R1 was introgressed from short-day S. demissum into potato plants together with some gene(s) conferring late transition to flowering.
Planta, 1977
The sequential appearance of a specific group of embryonal antigens (EA), presumably globulins, w... more The sequential appearance of a specific group of embryonal antigens (EA), presumably globulins, was demonstrated in developing maize (Zea mays L.) caryopses using a double immunodiffusion test with absorption of common antigens. Cross immunoelectrophoresis was employed to follow the differential pattern of EA accumulation in the growing scutellum and embryonic axis. The transient nature of two predominant EA seems to indicate their role as specific protein reserves of embryonal tissues. Another presumably organ-specific EA was maintained in callus obtained from a 28-day-old culture of scutellum isolated from the mature non-germinated caryopsis.
Biochemie und Physiologie der Pflanzen, 1982
ABSTRACT Erythrocyte-binding proteins (lectins) were found in the apical meristem, the stele and ... more ABSTRACT Erythrocyte-binding proteins (lectins) were found in the apical meristem, the stele and the cortex with epidermis of the primary root of maize seedlings, and some of these lectins were present only in the tissues possessing the vascular elements. We presumed that these vascular-specific lectins could be related to the stele-specific antigens reported previously (Khavkin et al. 1980). In the electrointmnnodiffusion test with polyvalent antisera against the stele or the cortex proteins, proteins solubilized from human erythrocytes preagglutinated with the stele extract produced two common antigens and three stele-specific precipitin bands. To investigate these lectins in more detail, the antiserum was raised in rabbits against stelar proteins immobilized on the rabbit erythrocyte ghosts. This antiserum precipitated up to five antigens in one-dimensional immunoelectropherograms of the root extracts, including the vascular-specific lectins absent from the cortex. One of the stele-specific ectins was apparently a precipitin-like lectin interacting with non-immune serum proteins.
Russian Journal of Plant Physiology, 2000
Russian Journal of Plant Physiology, 2005
The arabidopsis gene LEAFY controls the induction of flowering and maintenance of the floral meri... more The arabidopsis gene LEAFY controls the induction of flowering and maintenance of the floral meristem identity. By comparing the primary structure of LEAFY and its homologs in other Brassicaceae species and beyond this family, we singled out four clusters corresponding to three systematically remote families of angiosperms, Brassicaceae, Solanaceae, and Poaceae, and to gymnosperms. Both structural and functional distinctions of LEAFY homologs from their arabidopsis prototype expanded in the range Brassicaceae-Solanaceae-Poaceae. A LEAFY homolog from B. juncea cloned in our laboratory was used as a hybridization probe to analyze the restriction fragment length polymorphism in six Brassica species comprising diploid (AA, BB, and CC) and allotetraploid (AABB, AACC, and BBCC) genomes. In this way we recognized LEAFY fragments specific of genomes A, B, and C; in contrast, the variations of the length and structure of the LEAFY intron 2 were not genome-specific. LEAFY polymorphism in the Brassica accessions comprising genome B was related to their geographic origin and apparently to the adaptation to day length.
Zeitschrift für Pflanzenphysiologie, 1978
Antigens and isozyme spectra characteristic of epigenotypes of specialized cells in intact plants... more Antigens and isozyme spectra characteristic of epigenotypes of specialized cells in intact plants were preserved in successive subcultures of the four types of callus tissues derived from mesocotyls and scutella of inbreds CI03 and W155 and their crosses.
Zeitschrift für Pflanzenphysiologie, 1979
A shake flask culture was established from maize (hybrid C 103 X W 155) callus initiated at nodal... more A shake flask culture was established from maize (hybrid C 103 X W 155) callus initiated at nodal and internodal stem segments and maintained in the medium containing the inorganic constituents of MURASHIGE and SKOOG (1962), 20 gil sucrose, 0.4 mgll thiamine, 1 gil casein hydrolysate and 1 mgll 2,4-D. Small aggregates that lacked inner organization were predominant in the suspension. Cells were of a parenchyma-type, mostly round-shaped, and their sizes were distributed in log-normal pattern (the mean diameter 40-50 ft). The exponential growth in the batch-grown culture lasted to 4th-5th day, the transition to the stationary phase took place during the 5th-6th day, and after the 9th day a considerable degradation was observed in the culture. Doubling periods for the cell number, and dry matter and protein during the exponential growth were about 28 h at 26 DC, and the total increase was at least IS-fold during the 7-day subculture cycle. Respiration rate on the fresh weight, protein and cell basis was constant from the 1st to the 6th day and then declined 2-fold, however, it was completely restored during the first 12 h after cell transfer to the fresh medium.
Planta, 1980
A group of antigenically distinct proteins characteristic for the tissue complex of the vascular ... more A group of antigenically distinct proteins characteristic for the tissue complex of the vascular cylinders was found in maize (Zea mays L.) seedlings using an immunofiltration technique. Specific stelar antigens present in the fully developed stele (vascular cylinder) of the primary root were also found in steles extracted from adventitious roots and from the mesocotyl but were absent, within the limits of sensitivity of the immunodiffusion tests employed, in root cortex and epidermis. Some of the stelar antigens were also evident in the meristem of the primary root and were present in traces in the scutellum, the mesocotyl node, and the primary leaves plus coleoptile. The specific stelar antigens could be traced in 13-and 15-day-old developing embryos and were definitely expressed by the 21st day after pollination. Several stelar-specific antigens were found in embryo-derived callus tissues and in stem-derived cells maintained in serial suspension culture. Higher resolution of the stelar antigens by a modified technique of crossed immunoelectrophoresis was used to demonstrate several minor stelar antigens that were presumably characteristic exclusively of the completely differentiated stele. This technique along with sequential immunoprecipitation of labelled proteins provided a semiquantitative estimate of the specific stelar antigens in the meristem and the stele of the primary root, and in suspension-cultured cells which were devoid of noticeable signs of vascular differentiation.
Russian Journal of Plant Physiology, 1999
Solanum bulbocastanum comprising a CC-NBS-LRR gene RB/Rpi-blb1 confers broad-spectrum resistance ... more Solanum bulbocastanum comprising a CC-NBS-LRR gene RB/Rpi-blb1 confers broad-spectrum resistance to Phytophthora infestans and is currently employed in potato breeding for durable late blight (LB) resistance. Genomes of several Solanum species were reported to contain RB homologues with confirmed broad-spectrum defence function. With the discovery that novel P. infestans races break LB resistance mediated by the genes of broad-spectrum specificity, pyramiding several RBlike genes from various Solanum species in a single potato cultivar seems a promising approach to durable LB resistance. Here we report early evidence on RB-like sequences in the wide range of Solanum species section Petota. The panel of Solanum species was screened with three RBrelated PCR markers. RB-like sequences were found in every tested Solanum accession suggesting universal distribution of RB structural homologues among Solanum genomes, while the marker RB-629 corresponding to the RB gene was found in 14 speci...
SUMMARY Allelic diversity of the IpiO gene encoding the effector recognised by Rpi‐blb1/RB gene f... more SUMMARY Allelic diversity of the IpiO gene encoding the effector recognised by Rpi‐blb1/RB gene from S. bulbocastanum and S. stoloniferum was assessed by PCR screening and sequencing in 15 races (1, 3, 4, 10, 11, 1.2, 1.3, 1.4, 2.4, 3.4, 1.11, 1.2.3, 1.2.4, 1.3.4, and 1.2.3.4) maintained in the collection of the Institute of Phytopathology. IpiO locus was present in all these isolates. Phylogenetic analysis demonstrated that IpiO alleles from races 3 and 4 belonged to the classes I and II recognised by R pi-blb 1/RB. It follows that these “simple” races exhibit simple virulence patterns on potato comprising only R1-R11 genes from S. demissum . Growing interest in exploiting exotic Solanum germplasm in potato breeding is an incentive to reconsider and unambiguously redefine the terms “simple” and “complex” race. KEYWORDS Phytophthora infestans , Solanum , potato, late blight resistance, effectors, simple races INTRODUCTION Approaches to describe the diversity of P. infestans Various ...
Sel'skokhozyaistvennaya Biologiya, Jun 1, 2016
The authors are grateful to the Center for Collective Use of Equipment «Biotechnology» (All-Russi... more The authors are grateful to the Center for Collective Use of Equipment «Biotechnology» (All-Russian Research Institute of Agricultural Biotechnology) for sequencing Avr clones and separating SSR amplicons by capillary electrophoresis. Supported by the grants from Russian Foundation for Basic Research (projects 14-04-31613а and 16-04-00098) and the Ministry of Education and Science of the Russian Federation (project RFMEFI62114X0003). The phytopathological evaluation of Phytophthora infestans isolates was performed as part of the State Commission 0598-2015-0018.
Breeding potato for durable resistance to late blight (LB) greatly benefits from expanding the re... more Breeding potato for durable resistance to late blight (LB) greatly benefits from expanding the resource of resistance genes provided by wild Solanum species. SCAR markers for race-specific CC-NBS-LRR resistance genes (R genes) have been employed to screen Solanum accessions representing six series of section Petota. Structural homologues of particular genes were found in many taxonomically distant species and widely differed in their distribution patterns. Such evidence suggests that the R-gene structures evolved before the divergence of genomes A and B and preceded Solanum speciation.