CN117716996A - Crop breeding method for cultivating heterotrophic tetraploid - Google Patents
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Abstract
The invention relates to the technical field of crop genetic breeding, in particular to a crop breeding method for breeding heterotrophic tetraploids. The specific technical scheme is as follows: a crop breeding method for cultivating the heterotrophic tetraploid includes such steps as hybridizing the different diploid parents with homozygous genotype to obtain the first filial generation, and doubling the chromosome of the first filial generation to obtain the new tetraploid variety. The crop breeding method for cultivating the heterotrophic tetraploid provided by the invention organically combines the potential heterosis and polyploid advantage of the plant and gives full play to the heterosis, thereby cultivating a breakthrough high-yield high-quality new variety.
Description
Technical Field
The invention relates to the technical field of crop genetic breeding, in particular to a crop breeding method for breeding heterotrophic tetraploids.
Background
Plants can be classified into haplotypes (one chromosome group), diploids (two chromosome groups), polyploids (three and more chromosome groups), and aneuploidies (chromosome numbers are not the whole number of one chromosome) according to the number of chromosome groups of their cells.
Most plants in nature are diploid, with 2 sets of chromosomes, one from the female parent and one from the male parent. In diploid plants, there is a part of self-pollination, i.e. 1 plant provides both sperm and ovum, the chromosome set of which is identical to that of the ovum, and this plant belongs to a "genotype homozygous" diploid, e.g. soybean, rice. The seeds of the plant can be planted to obtain the same plant as the original plant.
In addition to self-pollination, another part of the plants in the diploid plant is cross-pollinated, i.e. 1 plant provides sperm and another 1 plant provides ovum, the chromosome set of which is not exactly the same as the chromosome set of the ovum, and this plant belongs to the "genotype heterozygous" diploid, e.g. orchid, litchi. Seeds of these genotype-heterozygous plants were sown, and plants having the same traits as those of the original parent plants could not be obtained.
Heterosis is a conceptual term of breeding that refers to the situation where a progeny plant has some advantage over the parent in crossing between two dissimilar plants. Typically, the advantage of crossing offspring of two different homozygous genotypes is more pronounced, while the advantage of crossing offspring between two different genotype heterozygous parents is weaker, or even completely less advantageous. Based on this situation, in order to allow the crossing of plants heterozygous for the genotype to have heterosis, it is necessary to transform such plants heterozygous for the genotype into plants homozygous for the genotype before crossing. This strategy is currently used for more heterosis breeding, e.g., the hybrid generation maize, hybrid generation tomato and most melons currently in production are produced (seeded) in this way.
To transform heterozygous genotypes into homozygous genotypes, the conventional practice is to repeatedly backcross and select for multiple generations. This method is time-consuming and labor-consuming, and is often prone to accidents in the middle, leading to waste of work. Compared with the time-consuming and labor-consuming multi-generation backcross method, the anther culture technology can achieve the purpose of purifying the heterozygous genotype in no more than half a year through one experimental operation. However, anther culture techniques have been successful for only a small fraction of plants, such as the families oryza, solanaceae and asteraceae.
Crop breeding is also commonly known as artificial mutation breeding in addition to the above cross breeding, while polyploid breeding belongs to one of the mutation breeding. As described above, since polyploids have 1 set (triploid), 2 sets (tetraploid) or even more than 2 sets of chromosomes relative to diploids, even if the genes in the set of chromosomes are identical, they are different from diploids to some extent due to the effect of the gene dose. At present, many researches show that polyploids are expected to have great changes in plant type (morphology), resistance, generation and accumulation of secondary metabolites and the like, and a few examples prove that polyploids have various advantages. In addition, many crops grown in large areas are now natural polyploids, such as bananas, sugarcanes, potatoes, strawberries, and the like.
The usual strategy for polyploid breeding is to use chemical agents to prevent normal cell division, so that the number of chromosomes in the cells is multiplied, thereby exerting the advantages of polyploid in certain aspects and improving the yield and/or quality. However, since polyploid seed setting rate is low, it is difficult to apply to plant types such as rice and soybean for seed harvest, and thus the range of application is severely limited.
The common practice in polyploid breeding is to treat the side and top shoots of the seed embryo and plant stem, while the most commonly used chemical agent is colchicine. Since the apical growth cone of these shoots is composed of many cells which differ in state due to subtle differences in the location where they are located, it is difficult to achieve the effect of doubling the chromosomes of all cells by chemical treatment, i.e., it is often only possible to obtain incompletely doubled "chimeras". The chimeric body is extremely unstable in the proportion state of cells with different ploidy due to different division periods of two ploidy cells, and the proportion of doubled cells is gradually reduced along with the growth of plants due to relatively longer cell period, so that the polyploid effect is finally disappeared.
Crop breeding, except for the most original natural mutation selection, the transgene which is only developed in recent years and molecular biological breeding by applying a gene editing technology, the most main breeding mode of modern breeding is hybrid breeding for creating mutation and hybrid breeding among genotype homozygous parents by utilizing heterosis. As described above, polyploid breeding works are relatively few because of being affected by the problems of low setting rate and great technical difficulty in achieving complete doubling.
Disclosure of Invention
Aiming at the defects of the prior art, the invention provides a crop breeding method for cultivating the heterotic tetraploid, which organically combines the potential heterosis and polyploid advantage of plants and fully plays the role, thereby cultivating a breakthrough high-yield high-quality new variety.
In order to achieve the above purpose, the invention is realized by the following technical scheme:
the invention discloses a crop breeding method for cultivating heterotrophic tetraploids, which is characterized in that hybrid generation is obtained by crossing different diploid parents with homozygous genotypes, and a new tetraploid variety is obtained by doubling the chromosome of the hybrid generation.
Preferably, anther culture is carried out by utilizing a diploid material heterozygous with a natural genotype to obtain homozygous heterozygous genotype diploid, then heterozygous genotype homozygous diploid is adopted in a mode of hybridization among different varieties to obtain genotype heterozygous hybrid generation, and then tissue culture is carried out.
Preferably, the hybrid generation is cultivated, the material is obtained from the offspring plant of the diploid parent hybridization combination which shows hybrid vigour, the tissue culture is carried out, the chromosome doubling is induced and the tetraploid plant regeneration is induced.
Preferably, vegetative organ tissue including, but not limited to, young leaves, young stems, young flower stems, and not including anther-containing inflorescences, is taken from plants as a culture material.
Preferably, the fully multiplied tetraploid plants are tissue-cloned so that each hybrid progeny line forms a population of a number that can be compared for field planting.
Preferably, the cloning is performed by a lateral bud proliferation method, and the cloning is performed by an adventitious bud proliferation method which is easy to generate mutation is avoided.
The invention has the following beneficial effects:
1. the invention organically combines the polyploid breeding strategy and the heterosis breeding which are commonly applied less, thereby obtaining the effect of overlapping the polyploid advantage on the basis of the heterosis, and further cultivating a new breakthrough crop variety.
2. On the basis of successful hybrid vigor breeding, the invention adds a technical link for inducing chromosome doubling, so that a new variety cultivated by the method adds a new advantage, namely polyploid advantage. It is generally thought that polyploid is expected to increase the yield by 30 to 50% due to the double increase in the number of genes, and that improvement or change in quality, stress resistance, and the like is also possible to some extent, relative to the original diploid variety.
Drawings
FIG. 1 is an inflorescence map of anther culture material;
FIG. 2 is an anther culture and callus map;
FIG. 3 is a drawing of adventitious buds regenerated from callus;
FIG. 4 is a chromosome map of haploids;
FIG. 5 is a haploid (right) and diploid plot of plants;
FIG. 6 is a graph of adventitious buds regenerated from a leaf explant;
FIG. 7 is a diagram of adventitious buds regenerated from a petiole explant;
FIG. 8 is a graph of root explant regenerated adventitious buds;
FIG. 9 is a diagram of a whole plant with adventitious bud rooting;
FIG. 10 is a colchicine doubling treatment diagram;
FIG. 11 is a diploid and tetraploid tissue culture Miao Tu;
FIG. 12 is a diagram of diploid and tetraploid chromosomes;
FIG. 13 is a clone diagram of lateral bud proliferation mode;
FIG. 14 is a diagram of a hybrid polyploid plant-bud phase;
FIG. 15 is a flowering phase diagram of a heteropolyploid plant.
Detailed Description
The following description of the embodiments of the present invention will be made clearly and completely with reference to the accompanying drawings, in which it is apparent that the embodiments described are only some embodiments of the present invention, but not all embodiments. All other embodiments, which can be made by those skilled in the art based on the embodiments of the invention without making any inventive effort, are intended to be within the scope of the invention.
The technical means used in the examples are conventional means well known to those skilled in the art unless otherwise indicated.
The invention discloses a crop breeding method for cultivating heterotrophic tetraploid, which comprises the following steps:
(1) For the natural genotype heterozygous diploid plant, the anther culture is carried out by directly taking materials. Haploids are obtained by anther culture, and chromosome doubling treatment is carried out on the haploids to obtain doubled haploids, namely, genotype homozygous diploids.
(2) Planting the double haploid plant regenerated by anther culture, and hybridizing (crossing with each other in the positive and negative directions) with the double haploid plant regenerated by anther culture to obtain the first filial generation seeds of each hybridization combination.
(3) Sowing and planting the seed fruiting body plants, comprehensively evaluating according to the performance of each hybrid combination offspring real plant group (for example, 10-20 seeds are obtained by each hybrid combination, 10-20 plant groups are formed by sowing and planting), and screening and determining the hybrid combination offspring with the most hybrid vigor.
(4) And respectively taking materials (diploid tissues such as tender leaves and the like) from hybrid combination offspring plants with obvious hybrid vigor, performing in-vitro culture, inducing chromosome doubling and tetraploid plant regeneration, and eliminating the chimera with incomplete chromosome doubling.
(5) And (3) carrying out seedling cloning in a lateral bud proliferation mode on each tetraploid regenerated plant with the completely doubled cell chromosome, and cloning 10-20 viable tissue culture seedlings from each plant.
(6) The tissue culture seedlings are planted, comprehensive comparison of all important characters is carried out, and a breakthrough excellent new variety with hybrid vigor and polyploid vigor is selected and cultivated.
The invention is further illustrated below in conjunction with specific examples.
The invention is further described in detail with respect to the cultivation of the Echinacea heterotrophic tetraploid.
Echinacea purpurea is an internationally known important medicinal crop, and has remarkable antibacterial, antiinflammatory and immunity enhancing effects. From the plant perspective, echinacea is a diploid plant with natural heterozygous cross pollination genotype, and is difficult to inherit stably through seeds, and polyploid Echinacea can solve the problem well through a asexual cloning seedling mode. In addition, the natural diploid plant has high plant type, is particularly easy to lodge in weather and rain, and causes great failure. Through multiple transformation, the branches can be obviously thickened, the lodging resistance is improved, and the purpose of high and stable yield is realized.
The method for cultivating the purple coneflower heterotrophic tetraploid comprises the following specific steps:
step 1: inflorescences with proper development period (see figure 1) are directly taken from physiologically mature plants and used as materials for anther culture. By suitable developmental stage is meant that the microspore cells within the anther of the floret on the head are at a "single-core by-edge" stage, and that the microspores at this stage are susceptible to external conditions to change direction of development, cease to develop into pollen and continue to divide, forming callus and eventually potentially differentiating to regenerate haploid plants.
Step 2: inflorescences are subjected to low-temperature treatment before anther culture. The aim of the low temperature treatment is to change the physiological state of inflorescences, especially microspores, so that the inflorescences are easier to accept the induction of anther culture conditions, and the formation rate of callus is improved. The low-temperature treatment condition for the Echinacea purpurea is 4-5 ℃ for 3-4 days.
Step 3: anther culture induced callus formation (see FIG. 2). Sterilizing the inflorescences subjected to low-temperature treatment, tearing off small buds by using pointed forceps, and clamping anthers for culturing. The culture medium for anther culture is prepared by adding 0.3mg/L of benzyl adenine and 0.1mg/L of naphthylacetic acid on the basis of an N6 formula, and increasing the concentration of sucrose to 6%; the culture condition is light-proof culture, and the time is 35-40 days.
Step 4: inducing the callus to differentiate to form adventitious buds (see FIG. 3). The calli formed by anthers are transferred and inoculated on a culture medium for inducing the differentiation of adventitious buds by forceps for culture. The culture medium for inducing the callus to differentiate adventitious buds is prepared by adding 0.5mg/L of benzyladenine and 0.05mg/L of naphthylacetic acid (sucrose concentration returns to 3% of usual) based on MS formula; the culture condition is illumination culture, and the time is 30-35 days.
Step 5: inducing adventitious buds to root to form complete plants. Adventitious buds are excised from callus and shallowly inserted onto rooting medium. The culture medium for inducing adventitious buds to root is obtained by adding 0.05mg/L of naphthylacetic acid based on an MS formula; the culture condition is illumination culture, and the time is 30-35 days.
Step 6: the whole plant regenerated by anther culture is essentially identified. Based on experience, echinacea can be identified by simple morphological observation. 1) Identification of haploids: compared with a diploid plant which is regenerated by culturing a leaf explant of an anther donor plant, the plant which has thinner leaves, finer leaf stalks and thinner roots is a haploid. 2) Identification of genotype homozygous diploids (originating from microspores and doubling from spontaneous chromosomes): the plant is a genotype homozygous diploid which has a significant difference in a certain trait from a diploid plant regenerated from an anther donor plant by leaf explant culture. The chromosome of haploid is shown in figure 4, and the haploid and diploid are shown in figure 5.
Step 7: haploid chromosome doubling and adventitious bud regeneration are induced. Chromosome doubling of haploid echinacea can be achieved by cutting leaf stalks from plants to serve as explants, performing induction treatment on a culture medium added with colchicine for 80-120 mg/L for inducing adventitious bud regeneration for 8-12 days, and then transferring the culture medium to a normal culture medium for inducing adventitious bud regeneration for culturing for 40-50 days, so that genotype homozygous diploid adventitious buds doubled in chromosome can be obtained. The colchicine doubling treatment is shown in FIG. 10, and the germination conditions are shown in FIGS. 6 to 8.
Step 8: the adventitious buds described in the step 7 are induced to root, and complete plants are formed (see FIG. 9). The specific method of this step is the same as that described in step 5.
Step 9: cloning the homozygous diploid plants of each anther culture origin, and planting a certain number of obtained cloned seedlings. The cloning adopts a lateral bud proliferation mode, and the method comprises the following steps: taking the base part of the plant (namely, the part left after most of the petioles and leaves are cut off), and culturing the plant on an MS culture medium added with 0.5-0.8 mg/L of benzyl adenine and 0.05mg/L of naphthylacetic acid; the culture condition is illumination culture, and the time is 30-35 days. This operation is repeated 2 to 3 times until a certain number of shoots are obtained, and then the shoots are induced to root to form a complete plant.
Step 10: planting each clone strain and hybridizing the strains to obtain hybrid first-generation seeds of each combination.
Step 11: sowing and planting seed actual plant of the hybrid combinations, and then comprehensively evaluating according to the performance of each hybrid combination offspring actual plant group (for example, 10-20 seeds are obtained by each hybrid combination, 10-20 plant groups are formed by sowing and planting), and screening and determining the hybrid combination offspring with the most hybrid vigor.
Step 12: the method is characterized in that the hybrid vigour is remarkable, a certain number (for example, 4-10) of ideal hybrid combination offspring plants are obtained (diploid tissues such as tender leaves) to carry out in vitro culture, chromosome doubling and tetraploid plant regeneration are induced (the method is the same as the step 7), and the chimera with incomplete chromosome doubling is eliminated.
Step 13: seedling cloning (plant morphology is shown in fig. 13) in a lateral bud proliferation mode is carried out on each confirmed fully doubled tetraploid regenerated plant, and 10-20 viable tissue culture seedlings are cloned from each plant. The diploid and tetraploid tissue culture seedling morphology and chromosomes are shown in fig. 11 and 12.
Step 14: the tissue culture seedlings are planted, comprehensive comparison of all important characters is carried out, and a breakthrough excellent new variety with hybrid vigor and polyploid vigor is selected and cultivated. The bud and flowering phases of the obtained heteropolynt plant are shown in fig. 14 and 15.
The above embodiments are only illustrative of the preferred embodiments of the present invention and are not intended to limit the scope of the present invention, and various modifications and improvements made by those skilled in the art to the technical solutions of the present invention should fall within the protection scope defined by the claims of the present invention without departing from the design spirit of the present invention.
Claims (6)
1. A crop breeding method for cultivating heterotrophic tetraploids, which is characterized by comprising the following steps: the new tetraploid variety is obtained through crossing different diploid parents with homozygous genotype to obtain hybrid generation and chromosome doubling the hybrid generation.
2. A crop breeding method for breeding heterotetraploids as recited in claim 1, wherein: anther culture is carried out by utilizing a diploid material heterozygous with a natural genotype to obtain a homozygous heterozygous genotype diploid, then a mode of hybridization among different varieties is adopted to heterozygote the genotype homozygous diploid to obtain a genotype heterozygous hybrid generation, and then tissue culture is carried out.
3. A crop breeding method for breeding heterotetraploids according to claim 1 or 2, characterized by: the hybrid generation is cultivated, the materials are obtained from the filial generation plants of the diploid parent hybridization combination which shows hybrid vigour, the tissue culture is carried out, the chromosome doubling is induced, and the tetraploid plant regeneration is induced.
4. A crop breeding method for breeding heterotetraploids as recited in claim 3, wherein: vegetative organ tissue including, but not limited to, young leaves, young stems, young flower stems, and not including anther-containing inflorescences, is taken from plants as a culture material.
5. A crop breeding method for breeding heterotetraploids as recited in claim 1, wherein: tissue culture cloning was performed on fully doubled tetraploid plants.
6. The method for crop breeding for heterotetraploid according to claim 5, wherein: cloning was performed by lateral bud proliferation.
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