CN102260736A - Identification method for distant hybrid progenies of peony - Google Patents
Identification method for distant hybrid progenies of peony Download PDFInfo
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Abstract
The invention relates to an identification method for hybrid progenies of plants, in particular to an identification method for distant hybrid progenies of peony. The identification method provided by the invention comprises the steps of: identifying and screening the early forms of a hybrid seedling, identifying the female parent by chloroplast DNA (deoxyribonucleic acid), and identifying the male parent by SSR (simple sequence repeat) molecular markers. By combining the form and molecule, the hybrid progenies are identified, true and false hybrids are distinguished, the relation between the true hybrid seedling and the parents is studied and summarized, and the heritable external form indices are determined; and on the molecular genetic level, the reasonable genetic distance between the hybrids and the parents and the distance of genetic relationship are derived, and finally a reasonable identification standard system is established on the form and molecular genetic levels, thus performing the later breeding process effectively.
Description
Technical field
The present invention relates to a kind of plant hybrid generation authentication method, particularly relate to a kind of distant hybrid of peony and Chinese herbaceous peony offspring's authentication method.
Background technology
Paeonia comprises about 35 kinds, is divided into three groups at present, i.e. tree peony group (sect.Moutan DC), Chinese herbaceous peony group (sect.Paeonia) and North America Chinese herbaceous peony group (sect.Onaepia Lindl)." flower king " tree peony is the time-honored traditional famous flower of Chinese cultivated with " flower mutually " Chinese herbaceous peony, have the good reputation of " two is exhausted in spending ", and its beauty is always praised highly by people.Because the two differs greatly in characteristic aspect such as blade profile, flower type, florescence and habits, so though they all are Paeoniaceae Paeonia plants, woody tree peony belongs to the tree peony group, the Chinese herbaceous peony of draft belongs to the Chinese herbaceous peony group.People generally believe that the hybridization that these two groups of plants are asked is " impossible " in the quite a long time in history, but development along with breeding technique, this " impossible " become reality, and a wonderful work that competitively matches in excellence or beauty with peony---the Itoh hybrid has occurred.
Distant hybirdization between group, the pollination back obtains seed easily and sprouts into individual plant, a large amount of filial generations is that cross-breeding work excellent selected good strains in the field for seed and selected and final-period management has increased difficulty, so need set up a cover easy early stage identification system of hybrid rationally as early as possible, filial generation is identified, differentiate true and false hybrid, real hybrid seedling of research summary and father and mother relation originally, determine heritable extraneous morphological index, in the molecular genetic aspect, draw the due genetic distance of rational cross-fertilize seed and parent, the far and near degree of sibship, on form and two aspects of molecular genetic, combine and set up rational standard of perfection system,, continue to observe to be selected for meeting then keeping for the time being of this requirement, otherwise directly eliminate, the seed selection process in later stage can effectively be carried out, in order to avoid waste of manpower, material resources and financial resources, this is the work that must pay attention in the Paeonia breeding process.
Used hybrid identification means mainly contain in ornamental plant distant hybridization breeding process at present: morphology mark identification method, armful powder are learned research identification method, cytological marker identification method, biochemical marker identification method and molecular markers for identification method.For the reliability of qualification result, generally can not a kind of method of single selection identify, but two kinds or more of method is used simultaneously.Be labeled as the master with morphology, auxiliary other marking methods of using.
And for the evaluation work of relevant Paeonia distant hybrid progeny, domestic rarely have research, and only report has Zhang Dong, Hao Qing, Guan Kun etc. to carry out the work of this respect at present.
Zhang Dong etc. (2008) utilize the AFLP technology that 7 clones of Paeonia papaveracea and 22 filial generations of ' high noon ' hybridization generation are identified, analytical results to specific band shows, the probability that maternal specific band occurs in for individuality at the F1 of distant hybirdization is more than male parent, and most hybrids and maternal sibship are near with respect to male parent; Also having drawn filial generation by calculating of cross combination genetic similarity and UPGMA cluster analysis is the conclusion of maternal partially pattern of fusion hybrid.
Hao Qing etc. (2008) find the distant hybrids of a Chinese herbaceous peony and tree peony first in Changping, Beijing, with this cross-fertilize seed called after ' harmony '.This is that domestic first strain is bloomed ' her rattan hybrid ', observes and the SRAP molecule marker by it being carried out morphological characters, has proved that fully ' harmony ' is group intermolecular hybrid kind.
Guan Kun (2009) to tree peony meat floral disc subgroup tree peony be female parent and keratin floral disc subgroup tree peony be male parent carry out the distant hybirdization gained between subgroup filial generation totally 158 strains carry out the research of form aspect and RAPD molecule marker aspect.The result shows that the RAPD technology can be carried out the true and false hybrid identification of distant hybrid progeny between effective subgroup, but for the evaluation of the sibship cluster between further true hybrid variety later on, then may need more further to study.
Domestic research to group species hybrid sibship is few, traces it to its cause, and main is because domestic group species hybrid resource is few on the one hand, as everyone knows, the group species hybrid originates from Japan, develops in the U.S., cost an arm and a leg and rare, therefore caused the scientific research material shortage of domestic this respect.And it is few about the work sutdy of hybrid identification aspect, then may be because distant hybirdization at present also is faced with many problems such as parent's cross incompatibility, pollination back sterility, hybridization time is oversize, parent's flowering asynchronism, sibship is indeterminate between the parent, and the hybrid germination rate is low, perhaps can sprout but F1 is low etc. for fertility.Be subjected to the influence of these objective condition, limited the research work of China aspect the miracle of this Paeonia breeding circle of Paeonia group species hybrid.
Along with the continuous appearance of distant hybrids new variety and the continuous progress of breeding work, in recent years, more and more organize species hybrid and be introduced into of the effort of domestic, domestic many scientific research institutions by continuing, new progress has also been arranged aspect distant hybridization breeding.Add that present dna sequencing, molecule marker equimolecular biological means all are applied to the research of Paeonia each side gradually, only can be more and more deeply and comprehensively to the research of Paeonia group species hybrid sibship, to organize species hybrid early molecule authenticate technology route efficiently also be imperative and set up a cover.
Summary of the invention
In order to address the above problem, the object of the present invention is to provide a kind of distant hybrid of peony and Chinese herbaceous peony offspring's authentication method.
Distant hybrid of peony and Chinese herbaceous peony offspring's provided by the invention authentication method comprises the steps:
1) hybrid strain and hybrid generation are carried out the morphological index preliminary screening;
2) hybrid generation and the father and mother that step 1) is filtered out originally carry out the chloroplast gene order-checking respectively;
3) hybrid generation and the father and mother that step 1) is filtered out originally carry out the SSR molecule marker respectively.
Wherein, step 1) also comprises processings of encoding of the nonumeric class proterties in the morphological index that will measure, and polynary proterties is handled by unordered proterties, calculates the genetic distance originally with father and mother.
Wherein, step 2) comprise that also the chloroplast gene to obtaining carries out Phylogenetic Analysis, identifies the female parent of filial generation.Preferably a plurality of chloroplast genes are carried out combined analysis
Wherein, step 3) comprises that also the SSR mark that amplification is obtained checks order, and calculates allelotrope number and gene frequency, identifies the male parent of filial generation.
Described chloroplast gene is preferably petB-petD, rps16-trnQ or their specific fragment.The petB-petD gene fragment of preferably using CTATCGTCCRACCGTTACWGAGGCT (SEQ ID No.1) and CAAAYGGATAYGCAGGTTCACC (SEQ ID No.2) to amplify amplifies the rps16-trnQ gene fragment with CGTTGCTTTCTACCACATCG (SEQ ID No.3) and TTACTCGGAGGTTCGAATCC (SEQ ID No.4).
Described SSR mark is preferably one or more among Pdel02-2, Pdel05, Pdel07, Pdel20, Pdel22, Pdel29b, Pdel33, Pdel35, Jx02-2 and the Jx17.
The mode that combines by form and molecule, filial generation is identified, differentiate true and false hybrid, real hybrid seedling of research summary and father and mother relation is originally determined heritable extraneous morphological index, in the molecular genetic aspect, draw the due genetic distance of rational cross-fertilize seed and parent, the far and near degree of sibship finally combines on form and two aspects of molecular genetic and sets up rational standard of perfection system, and the seed selection process in later stage can effectively be carried out.
Description of drawings
Figure 1 shows that 2 parents and hybrid generation's blade.Wherein, A is a hybrid; B is a male parent; C is maternal.
Figure 2 shows that the principal coordinate analysis figure of 2 parents and 11 hybrid generation's phenotypic characters of not blooming.
Figure 3 shows that the principal coordinate analysis figure of 2 parents and 4 hybrid generation's phenotypic characters of blooming.
Figure 4 shows that UMPGA evolutionary tree based on two segmental 21 filial generations of chloroplast gene and 2 parents.
Figure 5 shows that parent and filial generation share allelic distribution plan.Wherein, N-H, N-P and N-M represent hybrid, male parent and maternal proprietary allelotrope respectively; N-PH and N-MH represent allelotrope and the allelotrope maternal and that hybrid is shared that male parent and hybrid are shared respectively.
Embodiment
Following examples are used to illustrate the present invention, but are not used for limiting the scope of the invention.
Hybrid seedling preparation method: the male parent bud was adopted back in the dew look phase,, be placed on the 24h that dries in the shade under drying, the no sunlight point-blank condition at the indoor flower pesticide of taking.Pat flower pesticide to pollen and shed fully, separate pollen and flower pesticide with sieve, the pollen that sifts out is collected in the clean bottle, storage is standby in the refrigerator of labelled being positioned over-20 ℃.The plant of selecting robust growth is as female parent, in the maternal bud windbell phase or reveal the look phase with tweezers or free-hand petal and stamen are pinched or torn, is enclosed within on the fresh idea and with paper clip with the template bagging sack is rolled tightly.Maternal bagging began pollination after 3-4 days when column cap begins secreting mucus.Pollinate and before 11 o'clock or after 17, carry out, repeat three days continuously, award altogether three times.Then 8, by the end of September, when female parent becomes carpel crab oil look and fine fisssure, take, be seeded in the seedbed after gathering, after planting regularly water, keep ground moistening.Regularly water, loosen the soil, uprooting weed and spray medicine.
From 2003-2007, carry out the distant hybirdization of tree peony and Chinese herbaceous peony according to above-mentioned method, female parent flies lotus for P.lactiflora ' powder cloud ', male parent is P. * golden Supreme Being of lemoinei Group ' ' (L ' Esperance).In the seedling of these acquisitions, according to the form preliminary screening, remove those obviously unsuccessful seedling of hybridization, select most probable wherein to hybridize successful 21 strain seedling (being numbered H2, H3, H4, H5, G1, G2, G3, G6, G7, G8, G9, G10, G11, G12, G14, G15, G16, G17, G18, G19, G20), wherein have 4 strain seedling to bloom.All vegetable materials comprise its parent, all plant in National forest park, vulture peak, Beijing, tree peony study base.
2008 and in May, 2009 are carried out the morphological characters investigation to this 21 strain filial generation, between this research is chosen and can be reflected in the comparative morphology kind and the proterties of product difference between species totally 34 above 23 parts of materials (male parent, female parent, 21 hybrids) are carried out morphology investigation and analysis.Comprising 5 of binary proterties, 4 of quantitative characters, multistate character is 25 in order.In the proterties quantizing process, the binary proterties is with " 0 ", " 1 " expression; Quantitative character such as blade length etc. are directly taken numeric coding, i.e. the serial number proterties; Multistate character such as pattern, flower type etc. are encoded to 0,1,2 successively in order in order ... n-1.Wherein do not bloom as yet because of most of hybrid seedling, the form investigation index is mainly based on other indexs except that flower portion organ.Only 2 parents and the 4 strains hybrid seedling of blooming has been carried out the investigation of whole morphological indexs, 11 strain seedling have only been investigated its leaf portion feature and habit, and other 6 strain seedling are still young, and morphological specificity is representative not enough, so investigate.The feature of 34 phenotypic characters, type and quantity coding see Table 1.
The feature of table 1 phenotypic character, type and quantity coding
Annotate: the N=quantitative character; B=binary proterties; The unordered proterties of D=
Morphological index to above-mentioned 15 hybrid seedlings and parent thereof is handled, the processing of encoding of wherein nonumeric class proterties, polynary proterties is all handled by unordered proterties, utilize Gower (1997) likeness coefficient to carry out the calculating of genetic distance, use MVSP 3.1 softwares, use principal coordinates method (PCoA) to carry out the quantitative classification statistical study of phenotypic characteristic, the main morphological characters difference of 17 individualities is embodied on the space ordering structure figure that is made of two principal constituents.
Space principal coordinate analysis result (Fig. 2 of male parent, female parent and 11 filial generation individual level of not blooming, PCo1 26.071%, PCo2 23.434%) show, male parent ' golden Supreme Being ' (" L ' Esperance ") can clearly distinguish with maternal " the powder cloud flies lotus ".The space principal coordinate analysis result of male parent, maternal and 4 filial generation individual level of blooming (Fig. 3, PCo1 36.929%, PCo2 29.334%) shows that also these 4 filial generations can be significantly and this differentiation of father and mother.Wherein, 11 filial generations that do not bloom have been investigated the whole proterties except that flower portion feature, and 4 filial generations that bloom have been investigated the whole proterties that comprise colored portion feature.
Important difference is present in spends height, annual shoot length, compound leaf length, flower footpath and leaflet number.In fact, when we screen seedling according to morphological specificity at first, mainly be exactly the leaf portion morphological specificity (Fig. 1) that is dependent on them.15 hybrid generations' leaf portion form is all originally obviously different with father and mother.Leaf occuping between Chinese herbaceous peony group and the tree peony group.Investigation finds that also most of hybrid generation's annual shoot is long, compound leaf is long, the leaflet number is also placed in the middle.4 strains bloom the hybrid seedling its to spend height also be between father and mother between this.
The amplification and the order-checking of embodiment 2 chloroplast genes
21 hybrid generations of embodiment 1 and 2 parents' young leaflet tablet behind silica dehydrator, is used day Plant Genome of root biotechnology company to extract test kit and extracts total DNA, press the test kit specification sheets and operate.
Select two chloroplast gene fragment petB-petD, rps16-trnQ checks order.The amplimer of petB-petD is shown in SEQ ID No.1 and 2; The amplimer of rps16-trnQ is shown in SEQ ID No.3 and 4.
Finish on the My of BIO-RAD company CyclerTM Thermal Cycler type PCR instrument polymerase chain reaction (PCR).Its amplification system is 25 μ l system: template 20-30ng, 10 * PCR buffer, 2.5 μ l, and dNTP 80mmol/L, primer 5umol/L, Taq enzyme 1U (pool, Beijing star bio tech ltd) adds distilled water to 25ul.Amplification program is:
94 ℃ of pre-sex change 2min;
94 ℃ of sex change 30sec;
54 ℃ of annealing 30sec;
72 ℃, 2min extends,
Repeat 34 circulations;
72 ℃ are extended 10min.
Amplified production detects through 1% agarose gel electrophoresis.The PCR product carries out sequencing reaction after adopting the PEG8000 purifying.
The sequencing reaction system is 10 μ l, and reaction system is as follows:
Template (i.e. Shang Mian amplified production) 50-100ng, 5 * BigDye buffer, 2.7 μ l, BigDye 0.4 μ l, primer (5pmol/L) 1.0 μ l add distilled water and supply 10 μ l.The sequencing reaction program: 94 ℃, 15 seconds, 50 ℃, 15 seconds, 60 ℃, 4 minutes, 30 circulations added 72 ℃ again and extended 30 minutes.
The sequencing reaction product after the use sodium-acetate carries out purifying, carries out on AB 3730XL automatic sequencer (Applied Biosystems).Two chloroplast genes have all used the two ends amplimer as sequencing primer.The sequencing primer of petB-petD is shown in SEQ ID No.1 and 2; The sequencing primer of rps16-trnQ is shown in SEQ ID No.3 and 4.
Utilize software Sequcher (Ver4.6) that institute's calling sequence is proofreaded and splice, utilize Se-Al editor (Ver.2.0), carry out the manual arrangement contraposition.To the equal weighting of bases all in the sequence that sequences and as unordered proterties, the room is as the 5th kind of feature (01) processing of encoding.To two segmental combined analysis of chloroplast gene, use MEGA4.1 software and carry out Phylogenetic Analysis, by the position of phylogenetic tree, identify the female parent of filial generation.
Obtain 46 sequences altogether, (petB-petD after rps16-trnQ) sequence is carried out contraposition arrangement and space coding respectively, carries out sequence and merges, and the sequence total length is 2799bp to two cpDNA.Detecting 50 variant sites altogether, is brief information site entirely, comprising 4 insertion/disappearances, is respectively 253bp, 31bp, 5bp and 6bp, 6 mononucleotides disappearance, other all be the base replacement.With the sequence construct phylogenetic tree after merging, find that the phylogenetic tree of building with MP method, NJ method and UPGMA method does not have difference substantially, choose a phylogenetic tree of UPGMA method structure according to bootstrap and analyze.
From Fig. 4, can very clearly see, on the chloroplast gene level, identify that hybrid and Chinese herbaceous peony ' the powder cloud flies lotus ' form one for 21, and obviously separate with ' the golden Supreme Being ' of tree peony group.Because chloroplast gene is a plasma inheritance, understand that from molecular level Shanghai Stock Exchange 21 are identified that hybrid and Chinese herbaceous peony ' the powder cloud flies lotus ' have nearer maternal sibship, Chinese herbaceous peony ' the powder cloud flies lotus ' is the female parent of cross hybrid seedling.
The amplification and the order-checking of embodiment 3SSR mark
From Yuan Junhui (its Ph D dissertation: the primer that Paeonia papaveracea and Yan'an tree peony origin research 2009.12) optimize 10 pairs of polymorphism height 14 pairs of primers that filter out, can be used for laboratory sample increase by (table 3) (SEQ ID No.5～SEQ ID No.24).
Table 3SSR primer
Upstream primer 5 ' end at every pair of primer carries out fluorescent mark, Pdel05, Pdel20, Pdel29b, Pdel33 FAM mark wherein, Pdel02-2, Pdel07, Pdel22, Pdel35 HEX mark, Jx02-2, Jx17 ROX mark.
Finish on the My of BIO-RAD company CyclerTM Thermal Cycler type PCR instrument polymerase chain reaction (PCR).Its amplification system is 20 μ l system: template 20-30ng, PCR Mix (Beijing hundred Tyke biotechnology companies) 10 μ l, and each 1 μ l of 5 μ mol/L primers adds distilled water 20 μ l.
Amplification program is:
94 ℃ of pre-sex change 2min;
94 ℃ of sex change 30sec;
Ta (48-54 ℃) 30sec that anneals;
72 ℃, 1min extends,
Repeat 30 circulations;
72 ℃ are extended 10min.
Amplified production detects through 1% agarose gel electrophoresis.The PCR product carries out sequencing reaction after adopting the PEG8000 purifying.
Mark GeneScan TM-600LIZ 0.4 μ l (Applied Biosystems in amplified production adds, USA), with Pdel33, Pdel22, Pdel02-2 and Jx17 mix, Pdel05, Pdel07 and Pdel20 mix, Pdel29b, Pdel35 and Jx02-2 mix, through high-temperature denatured (95 ℃, 2-5 minute), last AB 3730XL automatic sequencer (AppliedBiosystems) detects, utilize GeneMapper ver.4.0 (Applied Biosystems, USA) read genotype and allelotrope data automatically, because allelic correct interpretation is the basis of all work, all data peak figure have been carried out repeatedly manual check and correction, and part has been difficult to the data of interpretation and the material of pcr amplification failure has carried out repeated experiments.
Utilize Power-Marker ver.3.25 (Liu and Muse, 2005) or hand computation allelotrope and gene frequency data, and data are carried out statistical study.
Share and proprietary allelic distribution based on three groups, further the allelic source of the individual level of analysis bank species hybrid.Fig. 5 shows, identifies that at 21 in the sample, all sample has the allelotrope from maternal and male parent significantly simultaneously, also has its distinctive allelotrope simultaneously, more clearly shows the hybridization fact of identifying hybrid.
The two bonded result shows that Chinese herbaceous peony ' the powder cloud flies lotus ' is the female parent of 21 filial generations, and tree peony ' golden Supreme Being ' (' L ' Esperance ') is its male parent.
The above only is a preferred implementation of the present invention; should be pointed out that for those skilled in the art, under the prerequisite that does not break away from the technology of the present invention principle; can also make some improvements and modifications, these improvements and modifications also should be considered as protection scope of the present invention.
Claims (7)
1. a distant hybrid of peony and Chinese herbaceous peony offspring authentication method, it comprises the steps:
1) hybrid strain and hybrid generation are carried out morphological index mensuration, preliminary screening hybrid generation;
2) hybrid generation and the parent that step 1) is filtered out carries out the chloroplast gene order-checking respectively;
3) hybrid generation and the parent that step 1) is filtered out carries out the amplification of SSR molecule marker respectively.
2. authentication method according to claim 1 is characterized in that, step 1) also comprises processings of encoding of the nonumeric class proterties in the morphological index that will measure, and polynary proterties is handled by unordered proterties, calculates the genetic distance originally with father and mother.
3. authentication method according to claim 1 is characterized in that step 2) comprise that also the chloroplast gene to obtaining carries out Phylogenetic Analysis, identifies the female parent of filial generation.
4. authentication method according to claim 1 is characterized in that, step 3) comprises that also the SSR mark that amplification is obtained checks order, and calculates allelotrope number and gene frequency, identifies the male parent of filial generation.
5. according to each described authentication method of claim 1～4, it is characterized in that described chloroplast gene comprises petB-petD, rps16-trnQ or their specific fragment.
6. authentication method according to claim 5, it is characterized in that the specific fragment of the rps16-trnQ that described chloroplast gene goes out for the specific fragment of the petB-petD that goes out with the primer amplification shown in SEQ ID No.1 and the SEQ ID No.2 and/or with the primer amplification shown in SEQ ID No.3 and the SEQ ID No.4.
7. according to each described authentication method of claim 1～4, it is characterized in that described SSR is labeled as one or more among Pdel02-2, Pdel05, Pdel07, Pdel20, Pdel22, Pdel29b, Pdel33, Pdel35, Jx02-2 and the Jx17.
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| CN109486991A (en) * | 2018-12-06 | 2019-03-19 | 南京农业大学 | Identify molecular labeling primer composition and its application of pears and apple intergeneric conjugal transfer |
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| CN114464248A (en) * | 2022-04-12 | 2022-05-10 | 北京市农林科学院信息技术研究中心 | Method and system for calculating genetic relationship in breeding material family |
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| CN102719543A (en) * | 2012-06-25 | 2012-10-10 | 中国科学院植物研究所 | Method for identifying plant varieties by utilizing chemical molecular formulas of nucleotides |
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| CN105075850A (en) * | 2015-08-18 | 2015-11-25 | 大连民族大学 | New camellia oleifera high-yield cultivation molecular design method |
| CN105695610A (en) * | 2016-04-12 | 2016-06-22 | 中国科学院华南植物园 | Molecular identification method and identification primer for Dalbergia odorifera T. Chen and Dalbergia rimosa Roxb. |
| CN109486991A (en) * | 2018-12-06 | 2019-03-19 | 南京农业大学 | Identify molecular labeling primer composition and its application of pears and apple intergeneric conjugal transfer |
| CN109486991B (en) * | 2018-12-06 | 2021-11-30 | 南京农业大学 | Molecular marker primer composition for identifying intergeneric hybrid of pear and apple and application thereof |
| CN113403413A (en) * | 2021-04-01 | 2021-09-17 | 河南科技大学 | cPPSSR (cyclic shift keying) marker primer developed based on peony chloroplast genome sequence and application |
| CN113403413B (en) * | 2021-04-01 | 2022-11-01 | 河南科技大学 | cPPSSR (cyclic shift keying) marker primer developed based on peony chloroplast genome sequence and application |
| CN114464248A (en) * | 2022-04-12 | 2022-05-10 | 北京市农林科学院信息技术研究中心 | Method and system for calculating genetic relationship in breeding material family |
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