Nuclear genome size and karyotype analysis in Papaver for BAC library construction

The objective of the research carried out is study of the pathway of alkaloid production in Papaver species and cell lines, and integration data on physical mapping of newly developed marker DNA sequences with existing difference in expression of genes for key enzymes of alkaloid biosynthesis. This...

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Опубліковано в: :Біополімери і клітина
Дата:2005
Автори: Kyrylenko, T.K., Martynenko, O.I., Alkhimova, O.G.
Мова:Англійська
Опубліковано: 2005
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Цитувати:Nuclear genome size and karyotype analysis in Papaver for BAC library construction / T.K. Kyrylenko, O.I. Martynenko, O.G. Alkhimova // Биополимеры и клетка. — 2005. — Т. 21, № 2. — С. 145-150. — Бібліогр.: 14 назв. — англ.

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Digital Library of Periodicals of National Academy of Sciences of Ukraine
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author Kyrylenko, T.K.
Martynenko, O.I.
Alkhimova, O.G.
author_facet Kyrylenko, T.K.
Martynenko, O.I.
Alkhimova, O.G.
citation_txt Nuclear genome size and karyotype analysis in Papaver for BAC library construction / T.K. Kyrylenko, O.I. Martynenko, O.G. Alkhimova // Биополимеры и клетка. — 2005. — Т. 21, № 2. — С. 145-150. — Бібліогр.: 14 назв. — англ.
collection DSpace DC
container_title Біополімери і клітина
description The objective of the research carried out is study of the pathway of alkaloid production in Papaver species and cell lines, and integration data on physical mapping of newly developed marker DNA sequences with existing difference in expression of genes for key enzymes of alkaloid biosynthesis. This research requires the knowledge on genome structure and organization, and development of genomic resources for detailed characterization of opium poppy genome. The work is focused on the investigation of some features of genome organization of Papaver somniferum and related species, Papaver bracteatum and Papaver rhoeas. These characteristics are necessary for the construction of a BAC library which would be used as appropriate genomic resource for characterization of karyotype changes in poppy stocks with altered alkaloid biosynthesis pathway. Some of these stocks (cell lines) were generated and differed in the types of alkaloids they accumulated. Роботу присвячено вивченню шляхів біосинтезу алкалоїдів у видів Papaver та інтегруванню даних фізичного картування нових маркерних послідовностей ДНК з відмінностями в експресії генів ключових ферментів біосинтезу алкалоїдів. Таке дослідження потребує знання структури геному та його організації, а також передбачає наявність геномних ресурсів задля детального аналізу геному опійного маку. У даному повідомленні визначено розмір геному опійного маку та зроб­лено каріотипічний аналіз Papaver somniferum і його спорід­нених видів, P. bracteatum і P. rhoeas. Отримані характеристи­ки є необхідними для створення ВАС бібліотеки, яка слугує базисом для цитогенетичного картування та пошуку змін каріотипу в лініях маку, що відрізняються за шляхами біоси­нтезу алкалоїдів. Вже одержано декілька клітинних ліній, які накопичують різні типи алкалоїдів. The objective of the research carried out is study of the pathway of alkaloid production in Papaver species and cell lines, and integration data on physical mapping of newly developed marker DNA sequences with existing difference in expression of genes for key enzymes of alkaloid biosynthesis. This research requires the knowledge on genome structure and organization, and development of genomic resources for detailed characterization of opium poppy genome. The work is focused on the investigation of some features of genome organization of Papaver somniferum and related species, Papaver bracteatum and Papaver rhoeas. These characteristics are necessary for the construction of a BAC library which would be used as appropriate genomic resource for characterization of karyotype changes in poppy stocks with altered alkaloid biosynthesis pathway. Some of these stocks (cell lines) were generated and differed in the types of alkaloids they accumulated.
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fulltext ISSN 0233-7657. Біополімери і клітина. 2005. T. 21. № 2 Г Е Н О М І Й О Г О РЕГУЛЯЦІЯ Nuclear genome size and karyotype analysis in Papaver for ВАС library construction Т. K. Kyrylenko, О. I. Martynenko, O. G. Alkhimova Institute of Molecular Biology and Genetics, National Academy of Sciences of Ukraine 150 Zabolotnoho str., 03143 Kyiv, Ukraine E. mail: alkliimiv@hotmail.com The objective of the research carried out is study of the pathway of alkaloid production in Papaver species and cell lines, and integration data on physical mapping of newly developed marker DNA sequences with existing difference in expression of genes for key enzymes of alkaloid biosynthesis. This research requires the knowledge on genome structure and organization, and development of genomic resources for detailed characterization of opium poppy genome. The work is focused on the investigation of some features of genome organization of Papaver somniferum and related species, Papaver bracteatum and Papaver rhoeas. These characteristics are necessary for the construction of a BAC library which would be used as appropriate genomic resource for characterization of karyotype changes in poppy stocks with altered alkaloid biosynthesis pathway. Some of these stocks (cell lines) were generated and differed in the types of alkaloids they accumulated. Key words: Papaver, alkaloids, genome size, bacterial artificial chromosome library, FISH. Introduction. The data from several different alka­ loid-producing plants suggest that their biosynthesis and accumulation involve a highly regulated process that includes cell-, tissue-, development and envi­ ronment-specific controls [1, 2] . The evolution of alkaloid pathways together with their cellular com- partmentation appears to be closely associated with the primary reactions from which they have evolved. Opium poppy, Papaver somniferum, is cultivated for its alkaloid-rich latex. Tyrosine decarboxylase (TyDC) is the first enzyme in poppy alkaloid bio­ synthesis and is encoded by a small gene family. Members of this family are differentially expressed in organs of the plant and cultivated cells [3 ]. With the availability of an increasing number of genes involved in alkaloid biosynthesis, increasing efforts would be made to identify the regulators [4] associated with the development of specialized cell types which relate to alkaloid biosynthesis and accumulation. Gene loca- © T. K. KYRYLENKO, O. I. MARTYNENKO, O. G. ALKHIMOVA, 2005 lization and isolation require a detailed knowledge of genome structure. While genetic maps provide infor­ mation on relative order of molecular markers and genes along the chromosomes, physical mapping pro­ vides data on the physical position of DNA sequence within a genome. P. somniferum, P. bracteatum and P. rhoeas belong to one taxonomic section within genus Papaver and produce morphinan and bentho- phenanthridine type alkaloids which belong to dif­ ferent biosynthetic pathways. Large-insert DNA libraries are one of the key resources that facilitate gene isolation by positional cloning and the analysis of genome organization, structure and evolution. The easy handling and propagation of the clones make BACs (bacterial artificial chromosomes) an invaluable tool in genomic research, used for a variety of applications, including physical mapping and genome sequencing [5]. The production of full-length cDNA molecules from geno­ mic DNA libraries, representing genes of interest, is of paramount importance in basic plant biology re- 145 mailto:alkliimiv@hotmail.com K Y R Y L E N K O T . K. , M A R T Y N E N K O O . I . , A L K H I M O V A O . G search as well as plant biotechnology [6]. Estimation of genome size, karyotype parameters, testing restric­ tion endonucleases patterns are the necessary charac­ teristics for BAC library construction. This study was undertaken to determine nuclear genome size and genomic distribution of ribosomal DNA loci and two families of repetitive DNA sequen­ ces in Papaver species representing one taxonomic section within genus Papaver, with the aim to expand the number of species where these characteristics are known, and aid in the construction of a BAC library wich would be used as appropriate genomic resource for characterization of karyotype changes in poppy stocks with altered alkaloid biosynthesis pathway. Material and Methods. Plant material. For DNA extraction and chromosome slide preparations we used seedlings and roots of poppy species, P. som- niferum, P. bracteatum and P. rhoeas (accessions from Kyiv Central Botanic Garden collection). DNA was extracted from etiolated seedlings following Au- subel et al. [7]. Determination of genome size. According to Dolezel et al. [8 ], approximately 50 mg of midrib was cut from a poppy, P. somniferum, young leaf and transferred to a glass Petri dish. About 10 mg of a young leaf of maize (Zea mays cv. CE-777) with 2C = = 5.43 pg DNA was added and served as an internal reference standard. The tissues were chopped simul­ taneously in 1 ml of Otto I buffer (0.1 M citric acid, 0.5 % v/v Tween 20). Crude suspension of isolated nuclei was filtered through a 50 pm nylon mesh. Nuclei were then pelleted (300 g, 5 min), resus- pended in 200 fil Otto I and incubated for 1 h at room temperature. Finally, 600 fil Otto II buffer (0.4 M Na 2 HP0 4 ) , supplemented with 50 [tg/ml RNAse and 50 i K g / m l propidium iodide (PI), was added. Samples were analysed using Partec PAS flow cytometer («Partec GmbH», «Munster», Germany) equipped with 488-nm argon laser. The gain of the instrument was adjusted so that peak representing maize Gl nuclei appeared approximately on channel 100 on histogram of relative fluorescence intensity when using 512-channel scale. About 5,000 nuclei were analysed at rate 10—25 nuclei/s. Three plants were measured per accession. Analysis of each plant was repeated three times on different days. Nuclear DNA content was calculated from individual measurements. Fluorescence in situ hybridization (FISH). FISH probe for ribosomal DNA was obtained by labelling a pTa71 DNA clone containing 18S, 5.8S and 26S rRNA genes [9] with biotin-16-dUTP («Roche», Germany) by Nick Translation. Along with rDNA probe two new clones were used. These were made by sonicating total genomic DNA from P. somniferum (Danish flag accession) to an average length of 200 to 600 bp. The fragments were denatured at 100 °C in 0.1 M sodium phosphate buffer, pH 7.5, for 10 min and incubated at 60 °C to C 0t of 0.02. The samples were treated with SI nuclease. The resulting highly repeated double-stranded DNA sequences were clo­ ned into pUCJ8 at the PstI site. From several thousand clones, 200 were chosen randomly for dot hybridization with 3 2P-labelled total DNA from the representatives of four Papaver taxonomic sections (Meconella, Pilosa, MacranthalPapaver, Argemoni- dium). Two clones, pPs21 and pPs41 gave very strong signals with P. somniferum DNA and no visible signals with three other sections members. These clones were selected for further studies on in situ and southern hybridizations. Digoxigenin-labelled probes for these repeats were prepared using PCR with M13 direct and reverse primers and pPs21 and pPs41 clones as a templates. Metaphase spreads were prepared according to Alkhimova et al. [10]. The slides were treated with 100 mg/ml RNAse in a 2 x SSC solution at 37 °C for 1 h in a humid chamber, washed 3 x 5 min in 2 x SSC at room temperature. After two washes in 2 x SSC the slides were treated in 4 % paraformaldehyde for 10 min at room temperature, washed in 2 x SSC, dehydrated in ethanol series, and air dried. Prior to hybridization, the probes were mixed in a solution containing 50 % formamide, 10 % dextran sulphate, 0.12 % SDS in 2 x SSC and 50 ng/,ul salmon sperm DNA. 1 fil of probe in 30 fil hybridization mixture per slide was used. The hybridization mixture was dena­ tured at 70 °C for 10 min and incubated on ice for 10—15 min before being added to the preparations. The chromosomes together with the probes were denatured at 70 °C for 5 min and the hybridization was performed overnight at 37 °C in a humid cham­ ber. The slides were then washed in 2 x SSC at 42 °C and rinsed in a stringent washing solution of 20 % formamide in 0.1 x SSC at 42 °C for 10 min, followed by several washes in 2 x SSC and 4 x SSC (0.2 % Tween). The sites of digoxigenin- and biotin-labelled probe hybridization were detected using anti-digo- xigenin fluorescein («Roche») and streptavidin conju­ gated to Cy3 («Sigma», USA), respectively. Finally, the preparations were counterstained with DAPI (0.2 146 N U C L E A R G E N O M E S I Z E A N D K A R Y O T Y P E ANALYSIS IN P A P A V E R 120 -I 100 - | 8 0 ' If 6 0 ' -o I 40- 20 - 0 - 0 ( W G / m l ) and mounted in Vectashield antifade solution («Vector Laboratories*, USA). The preparations were evaluated using Olympus BX60 microscope equipped with optical filter sets appropriate for DAPI, fluorescein and Cy3 fluo­ rescence. The images of DAPI, fluorescein and Cy3 fluorescence were acquired separately with a b / w CCD camera, which was interfaced to a PC running the ISIS software («Metasystems», «Altlussheim», Germany) . The images were superimposed after con­ trast and background optimization. Results and Discussion. Estimation of genome size of Papaver somniferum. Flow cytometry is a rapid technique that allows accurate estimation of nuclear DNA content [11] . To determine nuclear DNA con­ tent in absolute units , fluorescence intensity of nuclei is compared with that of nuclei isolated from a species with known nuclear genome size. Fluorescence of Pl-stained nuclei were analysed using a Partec PAS II flow cytometer. Nuclei isolated from Z. mays cv. CE-777 with known nuclear genome size (2C = = 5.43 pg DNA) were used as internal s tandard to estimate nuclear DNA content of P. somniferum in absolute units. Small amounts of leaf tissues (stan­ dard and sample) were simultaneously chopped in buffer, supplemented with PI and RNAse. Suspension of nuclei was filtered through 50 p.m nylon and stored on ice prior to analysis . 2C DNA content of P. somniferum was calculated according to formula: 2C nuclear DNA content = 5.43 * G, peak mean of Papaver r t r = ~ — : r^r = 6.46 pg. G, peak mean of Zea Genome size, which represent one copy of nuc­ lear DNA equal to 1C, was further determined as: Genome size = = 2C nuclear DNA content (pg) . Q 9 7 8 . 1 Q 9 B P R L L ] Fig. 1 . Estimation of nuclear DNA content using flow cytometry. Si­ multaneous analysis of nuclei iso­ lated from Zea mays U) (2n - - 22) used as internal standard and from Papaver somniferum (2) The results of the study showed (Fig. 1) that the size of P. somniferum nuclear genome equal 6.46 pg is smaller than previously estimated [12]. Karyotype of P. somniferum and related species, P. bracteatum and P. rhoeas. A karyotype, which is characterized by the number and morphology of chromosomes, is an important characteristic of a species. Methods for chromosome preparation and in situ hybridization essentially followed Heslop-Harri- son et al. [13] . Root tips were fixed, partially digested with enzymes and cells were spread on slides. At least ten well-spread metaphase plates with similar degree of chromatin condensation were used to make chromosomal measurements . For constructing the karyotype, the chromosomes were arranged in order of decreasing size and increasing asymmetry (Fig. 2) . Localization of rDNA loci and repetitive DNA clones. Fluorescence in situ hybridization on P. som­ niferum chromosomes showed that pPs21 and pPs41 DNA sequences were distributed mostly uniformly along all chromosomes with gaps near centromeres and nucleolar organizer regions (data not shown). FISH with P. bracteatum, the representative of the same taxonomic group Macranthal Papaver, is shown in Fig. 3. DNA sequences have been shown to be dispersed over the P. bracteatum genome. It had substantially more copies of pPs41 dispersed along most chromosomes (Fig. 3, b) than pPs21 which gave less number of dots. Preparation of BAC library construction. BAC library is a source of DNA clones, which will be used as landmarks for chromosome identification, and also for the search of clones consisted of single copy sequences. An important step toward the structural analysis of a functional DNA domains is the const­ ruction of a large-insert libraries. Their inserts repre­ sent large DNA fragments that can be easily localized on mitotic chromosomes using FISH and allow selec- 147 5 yon c Fig. 2. Karyoidiogram of Papaver somniferum (a), P. bracteatum (b) and P. rhoeas (c) showing the length of individual chromosomes and centromere position. Distribution of r D N A loci revealed by in situ hybridization with pTa71 probe t ion of BAC clones with def ic iency in repetitive D N A sequences . T h i s approach requires determinat ion of restriction e n d o n u c l e a s e s su i table for library const ­ ruct ion, vector se lec t ion , appropriate ratio of high molecular weight D N A a n d c h o s e n vector, optimal l igation and transformation condi t ions . Testing conditions for different restriction en­ zymes. EcoRI, BamHI, or Hindlll, the restriction e n z y m e s most frequently u s e d in BAC cloning [5]. H i g h molecular weight D N A of P. somniferum, obtai­ n e d by purification of protoplasts from the seed l ings in e n z y m e solut ion fo l lowed by e m b e d d i n g on low- melt ing-point agarose , was d iges ted with the five restriction e n z y m e s , which can be ranged according to Fig. 3. In situ hybridization showing the localization of repetitive sequences, pPs21 (a) and pPs41 (b), derived from opium poppy, on chromosomes of Papaver bracteatum. Arrows mark NORs , arrow­ heads indicate major pPs21 sites at the terminal region of two chromosomes increase of recognit ion s i tes , in the nex t order: Kpnl, Xbal, BamHI, EcoRI, Hindlll. As Hindi 11 restriction fragments conta ined T y D C sequence (hybridizat ion data not s h o w n ) , and Hindlll d igest ion gave more fragments , this e n z y m e is c h o s e n for the construction of P. somniferum BAC library. Logistics. T h e p lnd igo B A C - 5 (Hind11Y-Cloning R e a d y ) vector has been chosen for the P. somniferum BAC library construct ion. T h e vector is provided in a «ready- to -use» state . It has been l inearized at the unique restriction e n z y m e recognit ion site (Hindlll), dephosphory la ted and rigorously tes ted for purity and recombinant c loning eff iciency (Epicentre) . C o m p e ­ tent Escherichia coli D H 1 0 B cel ls («Gibco BRL», USA) and Gibco BRL Cel l -Porator Sys tem («Life Techno log ies* , U S A ) have b e e n u s e d to get effective and rel iable result. T h e vector to D N A ratio has been calculated to be in proportion approximate ly 1:10. Conclusions drawn. T h e cultured cell l ines which accumulated alkaloids of only o n e type , morphinan ( thebaine) or benthophenanthr id ine (sanguinarine) have been generated from P. bracteatum (Fig. 4 ) . As the members of the T y D C g e n e family are dif feren­ tially expres sed in cel ls and organs accumulat ing predominant alkaloid type [ 1 ] , o n e can a s s u m e that different g e n e s of this family are «responsible» for the alkaloid type produced. Southern hybridizat ion with T y D C probe have revealed the different organization of restriction fragments in unorganized callus culture [3, 1 4 ] . T h e opium poppy T y D C g e n e family is composed of about 15 m e m b e r s that are divided into two subfamil ies with regard to the sequence identity [3 ]. T h e s e two groups d is t inguish in organ-specif ic express ion , namely , the m e m b e r s of the T y D C l - l i k e subgroup are e x p r e s s e d abundant ly in roots , whi le the members of the T y D C 2 - l i k e subgroup — in roots and 1 4 8 Fig. 4. Thin-layer chromatographic analysis of alkaloids in Papaver bracteatum cultured cell clones: 1—4 — primary cell cultures (up to 5 months in culture); EM, EM5, EMIO — morphogenous lines; CA — dedifferentiated callus. Authentic standards of sanguinarine (S), protopine (P), allocryptopine (A) and thebaine (F) were also run stems. The members of the P. somniferum TyDC gene family show also differences in the develop­ mental, tissue-specific, and inducible expression pat­ terns. This gene family can be used as a molecular marker to study the regulation and localization of alkaloid biosynthesis in the poppy species. Fluorescence in situ hybridization has been per­ formed to investigate the physical distribution of repetitive clones along the chromosomes of P. somni­ ferum and P. bracteatum species. Two new clones have been used, pPs21 and pPs41. In situ hybridi­ zation on P. somniferum chromosomes has shown that pPs21 and pPs41 DNA sequences are distributed mostly uniformly along all chromosomes with gaps near centromeres and nucleolar organizer regions (data not shown). These repetitive sequences have been shown to be dispersed over the P. bracteatum genome distinguishing by the copy numbers of their repeats (Fig. 3). For the first time, the localization of the ribosomal DNA on P. bracteatum chromosomes is shown. BAC library is a source of new DNA clones, which will be used as landmarks for the chromosome identification, and also for searching the clones con­ sisted of single copy sequences. An important step toward the structural analysis of functional DNA domains is the construction of large-insert libraries. N U C L E A R G E N O M E S I Z E A N D K A R Y O T Y P E ANALYSIS IN P A P A V E R The construction of the representative ВАС library requires the determination of the genome size to obtain the appropriate amount of clones. The results of the study show that the size of the P. somniferum nuclear genome equal to 6.46 pg is smaller than previously estimated. The genome size of P. somni­ ferum (In = 22) which represents one copy of nuclear DNA equal to 1C, has been determined as 3100 Mb. Based on this haploid genome size and average insert size of fragments around 150 kb that can be cloned efficiently in the ВАС vector [5], the library repre­ senting five genome equivalents would consist of 100000 clones. Using cDNA probes will allow the verification of the coverage. The results obtained will be used for the construction of the large insert size genomic library as a new molecular resource. Acknowledgements. We are grateful to Dr J. Dolezel (Institute of Experimental Botany, Olomouc, Czech Republic) for providing us with technical faci­ lities, and P. Suchankova for technical assistance. This work was supported by the National Aca­ demy of Sciences of Ukraine and the International Atomic Energy Agency (Research Contract No. 12235/RBF). T. K. Кириленко, О. 1. Мартиненко, О. Г. Алхімова Визначення розміру ядерного геному і каріотипічний аналіз видів Papaver для створення ВАС бібліотеки Резюме Роботу присвячено вивченню шляхів біосинтезу алкалоїдів у видів Papaver та інтегруванню даних фізичного картування нових маркерних послідовностей ДНК з відмінностями в експресії генів ключових ферментів біосинтезу алкалоїдів. Таке дослідження потребує знання структури геному та його організації, а також передбачає наявність геномних ресурсів задля детального аналізу геному опійного маку. У даному повідомленні визначено розмір геному опійного маку та зроб­ лено каріотипічний аналіз Papaver somniferum і його спорід­ нених видів, P. bracteatum і P. rhoeas. Отримані характеристи­ ки є необхідними для створення ВАС бібліотеки, яка слугує базисом для цитогенетичного картування та пошуку змін каріотипу в лініях маку, що відрізняються за шляхами біоси­ нтезу алкалоїдів. Вже одержано декілька клітинних ліній, які накопичують різні типи алкалоїдів. Ключові слова: Papaver, алкалоїди, розмір геному, ВАС бібліотека, флуоресцентна гібридизація in situ. Т. К Кириленко, Е. И. Мартиненко, Е. Г. Алхимова Определение размера ядерного генома и кариотипический анализ видов Papaver для создания ВАС библиотеки Резюме Робота посвящена изучению пути биосинтеза алкалоидов у видов Papaver и интегрированию данньїх по физическому кар- 1 4 9 KYRYLENKO Т. К., MARTYNENKO О. I., ALKHIMOVA О. G. тированию новых маркерных последовательностей ДНК с различиями в экспрессии генов ключевых ферментов биосинте­ за алкалоидов. Такое исследование нуждается в знании струк­ туры генома и его организации, а также предполагает наличие геномных ресурсов для детального анализа генома опийного мака. В данном сообщении определен размер генома опийного мака и сделан кариотипический анализ Papaver somniferum и его родственных видов, P. bracteatum и P. rhoeas. Полученные характеристики необходимы для создания ВАС библиотеки, предоставляющей базис для цитогенетического картирования и поиска изменений кариотипа у линий мака, различающихся путями биосинтеза алкалоидов. Уже выделены несколько кле­ точных линий, накапливающих разные типы алкалоидов. Ключевые слова: Papaver, алкалоиды, размер генома, ВАС библиотека, флуоресцентная гибридизация in situ. REFERENCES 1. Facchini P. J, De Luca V. Phloem-specific expression of tyrosine/dopa decarboxylase genes and the biosynthesis of isoquinoline alkaloids in opium poppy / / Plant Cell.—1995.— 7.—P. 1811—1821. 2. Bird D. A., Franceschi V. R., Facchini P. J. A tale of three cell types: alkaloid biosynthesis is localized to sieve elements in opium poppy / / Plant Cell.—2003.—15.—P. 2626—2635. 3. Facchini P. J., De Luca V. Differential and tissue specific expression of a gene family for tyrosine/dopa decarboxylase in opium poppy / / J. Biol. Chem.—1994.—269.—P. 26684— 26690. 4. De Luca V., St Pierre B. The cell and developmental biology of alkaloid biosynthesis / / Trends Plant Sci.—2000.—5— P. 168—173. 5. Gindullis F., Dechyeva D., Schmidt T. Construction and characterization of а ВАС library for the molecular dissection of a single wild beet centromere and sugar beet (Beta vulgaris) genome analysis / / Genome.—2001.—44.—P. 846—855. 6. Moller S. G., Chua N. H. Chemical regulated production of cDNAs from genomic DNA fragments in plants / / Plant J.—2002.—32.—P. 615—622. 7. Ausubel F. M., Brent R., Kingston R. E., Moore D. D., Seidman J. G., Smith J. A., Struhl К Current protocols in molecular biology.—New York: Greene/Wiley Int., 1987 — 418 p. 8. Dolezel J., Greilhuber J., Lucretti S., Meister A., Lysak M., Nardi L., Obermayer R. Plant genome size estimation by flow cytometry: inter-laboratory comparison / / Ann. Bot.—1998.— 82.—P. 17—26. 9. Gerlach W., Bedbrook J. R. Cloning and characterization of ribosomal RNA genes from wheat an barley / / Nucl. Acids Res.—1979.—7.—P. 1869—1885. 10. Alkhimova O. G, Heslop-Harrison J. S., Shchapova A. I., Vershinin A. V. Rye chromosome variability in wheat-rye addition and substitution lines / / Chromosome Res.—1999.— 7, N 3.—P. 205—212. 11. Dolezel J., Bartos J., Voglmayr H., Greilhuber J. Nuclear DNA content and genome size of trout and human / / Cytometry.— 2003.—51 A.—P. 127—128. 12. Srivastava S., Lavania U. C. Evolutionary DNA variation in Papaver II Genome.—1991.—34.—P. 763—768. 13. Heslop-Harrison J. S., Schwarzacher Т., Anamthawat-Jonsson К In situ hybridization with automated chromosome denatura- tion / / Technique.—1991.—3.—P. 109—116. 14. Kyrylenko Т. K, Martynenko О. I., Alkhimova O. G. Organ- specific expression of tyrosine/dopa genes and the biosynthesis of isoquinoline alkaloids in Papaver sp. / / Ukr. Biochem. J.—2002.—74.—P. 35. УДК 575:577.21.12 Надійшла до редакції 18.12.03 150
id nasplib_isofts_kiev_ua-123456789-155232
institution Digital Library of Periodicals of National Academy of Sciences of Ukraine
language English
last_indexed 2025-12-07T18:05:11Z
publishDate 2005
record_format dspace
spelling Kyrylenko, T.K.
Martynenko, O.I.
Alkhimova, O.G.
2019-06-16T12:23:27Z
2019-06-16T12:23:27Z
2005
Nuclear genome size and karyotype analysis in Papaver for BAC library construction / T.K. Kyrylenko, O.I. Martynenko, O.G. Alkhimova // Биополимеры и клетка. — 2005. — Т. 21, № 2. — С. 145-150. — Бібліогр.: 14 назв. — англ.
DOI: http://dx.doi.org/10.7124/bc.0006E5
https://nasplib.isofts.kiev.ua/handle/123456789/155232
575:577.21.12
The objective of the research carried out is study of the pathway of alkaloid production in Papaver species and cell lines, and integration data on physical mapping of newly developed marker DNA sequences with existing difference in expression of genes for key enzymes of alkaloid biosynthesis. This research requires the knowledge on genome structure and organization, and development of genomic resources for detailed characterization of opium poppy genome. The work is focused on the investigation of some features of genome organization of Papaver somniferum and related species, Papaver bracteatum and Papaver rhoeas. These characteristics are necessary for the construction of a BAC library which would be used as appropriate genomic resource for characterization of karyotype changes in poppy stocks with altered alkaloid biosynthesis pathway. Some of these stocks (cell lines) were generated and differed in the types of alkaloids they accumulated.
Роботу присвячено вивченню шляхів біосинтезу алкалоїдів у видів Papaver та інтегруванню даних фізичного картування нових маркерних послідовностей ДНК з відмінностями в експресії генів ключових ферментів біосинтезу алкалоїдів. Таке дослідження потребує знання структури геному та його організації, а також передбачає наявність геномних ресурсів задля детального аналізу геному опійного маку. У даному повідомленні визначено розмір геному опійного маку та зроб­лено каріотипічний аналіз Papaver somniferum і його спорід­нених видів, P. bracteatum і P. rhoeas. Отримані характеристи­ки є необхідними для створення ВАС бібліотеки, яка слугує базисом для цитогенетичного картування та пошуку змін каріотипу в лініях маку, що відрізняються за шляхами біоси­нтезу алкалоїдів. Вже одержано декілька клітинних ліній, які накопичують різні типи алкалоїдів.
The objective of the research carried out is study of the pathway of alkaloid production in Papaver species and cell lines, and integration data on physical mapping of newly developed marker DNA sequences with existing difference in expression of genes for key enzymes of alkaloid biosynthesis. This research requires the knowledge on genome structure and organization, and development of genomic resources for detailed characterization of opium poppy genome. The work is focused on the investigation of some features of genome organization of Papaver somniferum and related species, Papaver bracteatum and Papaver rhoeas. These characteristics are necessary for the construction of a BAC library which would be used as appropriate genomic resource for characterization of karyotype changes in poppy stocks with altered alkaloid biosynthesis pathway. Some of these stocks (cell lines) were generated and differed in the types of alkaloids they accumulated.
Acknowledgements. We are grateful to Dr J. Dolezel (Institute of Experimental Botany, Olomouc, Czech Republic) for providing us with technical facilities, and P. Suchankova for technical assistance. This work was supported by the National Academy of Sciences of Ukraine and the International Atomic Energy Agency (Research Contract No.
 12235/RBF).
en
Біополімери і клітина
Геном та його регуляція
Nuclear genome size and karyotype analysis in Papaver for BAC library construction
Визначення розміру ядерного геному і каріотипічний аналіз видів Papaver для створення ВАС бібліотек
Определение размера ядерного генома и кариотипический анализ видов Papaver для создания ВАС библиотеки
published earlier
spellingShingle Nuclear genome size and karyotype analysis in Papaver for BAC library construction
Kyrylenko, T.K.
Martynenko, O.I.
Alkhimova, O.G.
Геном та його регуляція
title Nuclear genome size and karyotype analysis in Papaver for BAC library construction
title_alt Визначення розміру ядерного геному і каріотипічний аналіз видів Papaver для створення ВАС бібліотек
Определение размера ядерного генома и кариотипический анализ видов Papaver для создания ВАС библиотеки
title_full Nuclear genome size and karyotype analysis in Papaver for BAC library construction
title_fullStr Nuclear genome size and karyotype analysis in Papaver for BAC library construction
title_full_unstemmed Nuclear genome size and karyotype analysis in Papaver for BAC library construction
title_short Nuclear genome size and karyotype analysis in Papaver for BAC library construction
title_sort nuclear genome size and karyotype analysis in papaver for bac library construction
topic Геном та його регуляція
topic_facet Геном та його регуляція
url https://nasplib.isofts.kiev.ua/handle/123456789/155232
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