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1.
Summary An attempt was carried out to produce trisomics of the wild tomato L. peruvianum, to define their essential features, and to detect relationships between trisomy and the expression of self-compatibility.Triploid-diploid crosses in L. peruvianum yielded nearly 40% aneuploids. Of these, 18% were single trisomics, and the rest had 2, 3 and 4 extra chromosomes. Almost all the trisomics occurred in crosses where the triploid was used as female parent. Vigour and fertility of trisomics were not much different from those of disomics, and morphologically they were very similar.The extra chromosome was identified in three self-compatible trisomic plants through somatic and pachytene chromosome morphology. One of these plants was trisomic for chromosome 1, while the other two were trisomic for chromosome 3. In these trisomics a positive correlation was found between chromosome length and trivalent formation, but no relationship between chromosome length and frequency of laggards was observed.A series of test-crosses revealed that the capacity of the trisomics to produce seed upon selfing always resulted from alterations of the incompatibility phenotype of the style and not from competitive interaction in the pollen. Progeny analyses showed that the self-compatibility features of the trisomics were not transmitted from one generation to the next. The implications of these findings are discussed.This work has been supported by a contract between the European Communities and the CNEN. This publication is contribution no. 1458 from the Biology Division of the European Communities and contribution no. 472 from the Divisione Applicazioni delle Radiazioni del CNEN.  相似文献   

2.
W Lange  T S Bock  G J Speckmann  J H Jong 《Génome》1993,36(2):261-267
Alien chromosome transmission through the female germ line as well as meiosis in pollen mother cells were studied in disomic and ditelosomic alien chromosome additions of beet. Beta vulgaris, carrying an extra pair of chromosomes or telosomes of B. procumbens or B. patellaris, respectively. The alien chromosomes carried genes for resistance to the beet cyst nematode, Heterodera schachtii, and screening for this resistance was used to select plants with or without the alien chromosomes. A great variation for alien chromosome transmission was recorded and plants carrying two extra alien chromosomes were recovered in the backcross progenies of the disomic or ditelosomic additions. However, in these progenies the average frequencies of plants without alien chromosomes (86%) did not clearly differ from that in similar progenies of the original monosomic or monotelosomic chromosome additions, indicating that doubling the number of the alien chromosome did not enlarge their transmission to the next generation. The alien chromosomes fully paired at pachytene and desynapsed again before diakinesis, indicating decreased chiasma formation. At second metaphase nearly 60% of the cells had one extra chromosome, and the remaining cells carried two or no extra chromosomes in about equal proportions. The tetrads looked fully normal. The expected relation between the average number of alien chromosomes in the germ cells and in the plants of the progenies did not show up, indicating a strong selection favouring the female gametes without alien chromosomes.  相似文献   

3.
Summary Trisomies of primary trisomic line B220 of Nicotiana sylvestris, which contain an extra chromosome shown to be a satellite chromosome, can be readily identified by their larger flower and leaf sizes. In seed-propagated species, the low transmission of the extra chromosome has prevented such plants from becoming agriculturally useful cultivars. In line B220, the transfer of the extra chromosome in 2n×2n+1 crosses was very low (13.5%), although n and n + 1 pollen grains were produced in equal quantities, as was confirmed by anther culture. This was due to the delayed development of n + 1 pollen grains, which are not at full maturity at the time of an thesis. The transfer of the extra chromosome in 2n×2n+1 crosses was increased by a 1 day delay in pollination and also by pollination of small pollen grains selected through nylon meshes. The delayed pollination increased the frequency of trisomics by 9%, whereas pollen selected by using 30 and 25 n nylon meshes induced an extremely high transfer of the extra chromosome, namely 51.9% and 70.4%, respectively. The observed frequencies of trisomics and tetrasomics in artificial selfing of 2n+1 plants with selected small pollen grains were lower than those expected from the data of reciprocal crosses between 2n and 2n+1 plants. This discrepancy seems to indicate a disadvantage of the n+1 pollen in fertilization due to the longer style of the trisomics relative to that of the diploids.  相似文献   

4.
Aneuploids have been identified cytomorphologically in progenies from triploid and diploid-triploid crosses. 30 trisomics, 2 double trisomics, 1 tetrasomic, and 2 higher aneuploids were obtained. Some of the aneuploids were found to be different from those reported earlier and higher aneuploids carried eight extra chromosomes. The plants with one extra chromosome occurred more frequently (67%) than the other aneuploid types. The changes in morphological traits such as a reduction in the amount of growth and the size of leaf etc. distinguished aneuploids from diploids. In the higher aneuploids the plant parts were highly exaggerated and pollen sterility was very high. The chromosomal counts in acetocarmine squashes confirmed the presence of extra chromosome(s). Aneuploids particularly trisomics were found to be promising and may lead to the production of commercially viable plants.  相似文献   

5.
Summary The karyotype of 82 regenerated plants from callus cultures of interspecific hybrids between cultivated barley (Hordeum vulgare L.) and seven polyploid wild barley species was examined by C-banding or Feulgen staining. The karyotypic changes observed in 46 plants included aneuploidy, double haploidy, amphidiploidy, deletions, inversions, extra C-bands, and extra euchromatic segments. Apparently, chromosome 5, 6, and 7 of H. vulgare were more frequently exposed to elimination or structural change than the other chromosomes of this species. Irradiation of calli seemed to enhance the occurrence of karyotypic variants.  相似文献   

6.
Summary Mesophyll protoplasts of the kanamycin-resistant nightshade, Atropa belladonna, were fused with mesophyll protoplasts of the phosphinothricin resistant-tobacco, Nicotiana tabacum. A total of 447 colonies resistant to both inhibitors was selected. Most of them regenerated shoots with morphology similar to one of the earlier obtained and described symmetric somatic hybrids Nicotiana + Atropa. However, three colonies (0.2%) regenerated vigorously growing tobacco-like shoots; they readily rooted, and after transfer to soil, developed into normal, fertile plants. Unlike their tobacco parental line, BarD, the obtained plants are resistant to kanamycin [they root normally in the presence of kanamycin (200 mg/1)] and possess activity of neomycin phosphotransferase (NPT II) with the same electrophoretic mobility as the one of the nightshade line. According to Southern blot hybridization analysis carried out with the use of radioactively labeled cloned fragments of the Citrus lemon ribosomal DNA repeat, as well as with Nicotiana plumbaginifolia genus-specific, interspersed repeat Inp, the kanamycin-resistant plants under investigation have only species-specific hybridizing bands from tobacco. Cytological analysis of the chromosome sets shows that plants of all three lines possess 48 large chromosomes similar to Nicotiana tabacum ones (2n = 48), and one small extra chromosome (chromosome fragment) similar to Atropa belladonna ones (2n = 72). Available data allow the conclusion that highly asymmetric, normal fertile somatic hybrids with a whole diploid Nicotiana tabacum genome and only part (not more than 2.8%) of an Atropa belladonna genome have been obtained without any pretreatment of a donor genome, although both these species are somatically congruent.  相似文献   

7.
Development and applications of a complete set of rice telotrisomics   总被引:8,自引:0,他引:8  
Cheng Z  Yan H  Yu H  Tang S  Jiang J  Gu M  Zhu L 《Genetics》2001,157(1):361-368
We previously isolated a complete set of primary trisomics along with many other aneuploids from triploid plants derived from an indica rice variety "Zhongxian 3037." About 30,000 progeny from these trisomic and aneuploid plants were grown each year from 1994 to 1999. The variants that differed morphologically from both the diploids and the original primary trisomics were collected for cytological identification. From these variants, a complete set of telotrisomics covering all 24 rice chromosome arms was obtained. The identities of the extra chromosomes were further confirmed by dosage analysis of the RFLP markers on extra chromosome arms. The telocentric nature of the extra chromosomes in these stocks was verified by fluorescence in situ hybridization (FISH) using a rice centromeric BAC clone as a marker probe. In general, the shorter the extra chromosome arm of a telotrisomic, the stronger the resemblance it bears to the diploid; the longer the extra chromosome arm, the stronger the resemblance to the corresponding primary trisomic. We demonstrated that DNA clones can be rapidly assigned to specific chromosome arms by dosage analysis with the telotrisomics. We also showed that telotrisomics are valuable tools for chromosome microdissection and for developing chromosome-specific DNA markers.  相似文献   

8.
Paternal inheritance of mitochondria in rapeseed (Brassica napus)   总被引:2,自引:0,他引:2  
Summary Transfer of a mitochondrially associated plasmid following sexual crosses in Brassica napus rapeseed suggested that paternal mitochondria were being transferred to the cytoplasm of the egg. To examine this possibility further, plants carrying the chloroplast (cp) marker of triazine resistance, but which had lost the plasmid associated with the mitochondria of this cytoplasm, were crossed as females to males carrying the polima cytoplasm. The males carried a nuclear fertility restorer gene on an extra chromosome to overcome the male sterility marker conferred by the mitochondria of this cytoplasm. Approximately 10% of the F1 progeny displayed the male sterility and flower morphology of the male parent. Mitochondrial (mt) DNA from the progeny showed the combined restriction patterns of both parents, but this rut heterogeneity did not continue into subsequent generations. All progeny retained the cp DNA restriction patterns of the maternal plant as well as resistance to the herbicide atrazine. To date, sexually mediated cybrid plants have shown no morphological abnormalities and have maintained their unique combination of cp and mt traits through several sexual generations.  相似文献   

9.
Summary Twelve distinct phenotypic groups of plants were isolated from nondisjunction progenies of 11 translocation heterozygote stocks. All the plants in these phenotypic groups originated in the light weight seed class. Five of the 12 phenotypic groups of plants have been verified as primary trisomics. They are all phenotypically distinguishable from each other and from disomics. One of the five primary trisomic groups, puckered leaf, was directly recovered as a primary trisomic from the original translocation heterozygote progenies. Three of the five trisomics — weak stem, dark green leaf, and convex leaf — originated first as tertiary trisomics. The related primary trisomics were isolated later from progenies of selfed tertiary trisomics. The fifth group, chlorotic leaf, originated at a low frequency among the progenies of three other trisomics: puckered leaf, convex leaf, and dark green leaf. The chlorotic leaf did not set seed under field conditions. The remaining four groups — puckered leaf, dark green leaf, convex leaf, and weak stem — are fertile, though sensitive to high temperature conditions. The transmission rate of the extra chromosome on selfing ranges from 28% to 41%. Physical identification of the extra chromosome has not been achieved for any of the five trisomic groups. Two trisomic groups, dark green leaf and convex leaf, have produced tetrasomics at low frequency. The phenotypes of these two tetrasomics are similar to the corresponding trisomics but more exaggerated.Fla. Agr. Expt. Stn. Journal Series No. 7137  相似文献   

10.
Ge XH  Li ZY 《Plant cell reports》2006,25(10):1075-1080
Abnormal meiosis and microspore development and related defective mutants have often been reported in plants and wide hybrids. Here extra divisions and nuclei fusions were observed to occur in microspore nuclei of partial hybrids between synthetic Brassica hexaploid (2n=54, AABBCC) and another crucifer Orychophragmus violaceus (2n=24). Abnormal spindle were formed and chromosomes were separated into several nuclei of variable sizes after bi-, or multi-polar divisions in the four cells of tetrads. As a consequence, more than eight mini-microspores of different sizes were produced by one tetrad. Genomic in situ hybridization results indicated that no chromosome replication occurred during such divisions. In some tetrads, the four nuclei were fused to form one large cell with increased chromosome number. The extra divisions or fusions appeared only in some flower buds of one plant, some anthers in the same buds, or even in individual cells of tetrads. The possible mechanisms behind these cytological phenomena are discussed.  相似文献   

11.
J F Golz  V Su  A E Clarke  E Newbigin 《Genetics》1999,152(3):1123-1135
Mutations affecting the self-incompatibility response of Nicotiana alata were generated by irradiation. Mutants in the M1 generation were selected on the basis of pollen tube growth through an otherwise incompatible pistil. Twelve of the 18 M1 plants obtained from the mutagenesis screen were self-compatible. Eleven self-compatible plants had mutations affecting only the pollen function of the S locus (pollen-part mutants). The remaining self-compatible plant had a mutation affecting only the style function of the S locus (style-part mutant). Cytological examination of the pollen-part mutant plants revealed that 8 had an extra chromosome (2n + 1) and 3 did not. The pollen-part mutation in 7 M1 plants was followed in a series of crosses. DNA blot analysis using probes for S-RNase genes (encoding the style function of the S locus) indicated that the pollen-part mutation was associated with an extra S allele in 4 M1 plants. In 3 of these plants, the extra S allele was located on the additional chromosome. There was no evidence of an extra S allele in the 3 remaining M1 plants. The breakdown of self-incompatibility in plants with an extra S allele is discussed with reference to current models of the molecular basis of self-incompatibility.  相似文献   

12.
Vasek , F. C. (U. California, Riverside.) Trisomic transmission in Clarkia unguiculata. Amer. Jour. Bot. 48(9): 829–833. 1961.—Seven primary trisomic plants derived from a triploid-diploid cross were self-pollinated. The 7 progenies included diploids and trisomics, the latter varying in frequency from 16 to 30%. In addition, 2 of the progenies included tetrasomic plants. Crosses were made between diploids and either trisomics or tetrasomics. The extra chromosome of 1 progeny was readily transmitted through the pollen of trisomic and tetrasomic plants. When a trisomic of the same progeny was used as a seed parent, only diploids and tetrasomics were found among the offspring, indicating a duplication of the extra chromosome. The extra chromosomes of other progenies were not transmitted through either pollen or eggs in controlled diploid-trisomic crosses but trisomics of these progenies were recovered after self-pollination. It is suggested that differential pollen-tube growth precluded transmission to diploid-trisomic hybrids and that under conditions of reduced pollen competition the extra chromosome normally would be transmitted through pollen. The extra chromosomes generally occur as univalents at metaphase and are ordinarily included in telophase nuclei.  相似文献   

13.
Summary Anther culture of secondary octoploid triticale (AABBDDRR) and F1 hybrids (AABBDDR) of octoploid triticale x common wheat crosses was carried out, and 96 pollen-derived plants were developed and studied cytologically. In addition to the 8 types of pollen-derived plants with the theoretically predicted chromosome number, plants with the chromosome constitutions of 2n = 38, 43, 45, 47, 74, and mixoploids were obtained. The haploids and the diploids had different distributions. The frequencies of plants with one and two (pairs of) rye chromosomes were extremely high, and anther culture may be an expeditious route for creating alien addition lines of distant hybrid F1s. Chromosome aberrations, including deletions, inversions, translocations, as well as isochromosomes and ring chromosomes, were observed in some plants. Abnormal meioses, such as chromosome non-disjunction, were also found. The reasons for the chromosome aberrations are discussed.  相似文献   

14.
Fras A  Maluszynska J 《Genetica》2004,121(2):145-154
Twelve callus lines of Arabidopsis thaliana were derived from four types of explants excised from diploid plants of two ecotypes (Columbia and Wilna) and autotetraploid plants of the Wilna ecotype. Cytogenetic analysis of the chromosome variation in particular callus lines was carried out for primary culture and callus during 5 months of culture. Ploidy levels of interphase nuclei were estimated by counting the number and size of chromocentres and nuclei of interphase cells. The first polyploid cells in all callus lines were observed during callogenesis. In primary culture the ploidy level ranged between 2 and 15x (10-75 chromosomes). The frequency of polyploid cells was higher in the 5-month old callus culture, but the ploidy level was the same. In the callus lines derived from autotetraploid plants, cells with reduced chromosome number appeared quite frequently along with diploid and polyploid cells.  相似文献   

15.
Aneuploids of sugi (Cryptomeria japonica) were found in the open-pollinated progenies of triploidplus tree clones. Seven trisomics and one hypotriploid were used to assign the chromosomes to the RFLP linkage groups constructed previously. The Southern blots containing their genomic DNA were hybridized with the labeled DNA clones corresponding to the loci in the linkage map. The additional dosage in autoradiographs showed that the cloned DNA fragment was located on the extra chromosome in the trisomics. On the other hand, the extra chromosome in two trisomics and the chromosome lacking the triplet in the hypotriploid were cytologically identified as chromosome 10 by consistent presence of a secondary constriction in the proximal region of its short arm. As a result, three linkage groups were assigned to their respective chromosomes, namely chromosome 10 and two other chromosomes.  相似文献   

16.
In cereals, interspecific and intergeneric hybridizations (wide crosses) which yield karyotypically stable hybrid plants have been used as starting points to widen the genetic base of a crop and to construct stocks for genetic analysis. Also, uniparental genome elimination in karyotypically unstable hybrids has been utilized for cereal haploid production. We have crossed hexaploid oat (2n=6x=42, Avena sativa L.) and maize (2n=2x=20, Zea mays L.) and recovered 90 progenies through embryo rescue. Fifty-two plants (58%) produced from oatxmaize hybridization were oat haploids (2n=3x=21) following maize chromosome elimination. Twenty-eight plants (31%) were found to be stable partial hybrids with 1–4 maize chromosomes in addition to a haploid set of 21 oat chromosomes (2n=21+1 to 2n=21+4). Ten of the ninety plants produced were found to be apparent chromosomal chimeras, where some tissues in a given plant contained maize chromosomes while other tissues did not, or else different tissues contained a different number of maize chromosomes. DNA restriction fragment length polymorphisms (RFLPs) were used to identify the maize chromosome(s) present in the various oat-maize progenies. Maize chromosomes 2, 3, 4, 5, 6, 7, 8, and 9 were detected in partial hybrids and chromosomal chimeras. Maize chromosomes 1 and 10 were not detected in the plants analyzed to-date. Furthermore, partial self-fertility, which is common in oat haploids, was also observed in some oat-maize hybrids. Upon selfing, partial hybrids with one or two maize chromosomes showed nearly complete transmission of the maize chromosome to give self-fertile maize-chromosome-addition oat plants. Fertile lines were recovered that contained an added maize chromosome or chromosome pair representing six of the ten maize chromosomes. Four independently derived disomic maize chromosome addition lines contained chromosome 4, one line carried chromosome 7, two lines had chromosome 9, one had chromosome 2, and one had chromosome 3. One maize chromosome-8 monosomic addition line was also identified. We also identified a double disomic addition line containing both maize chromosomes 4 and 7. This constitutes the first report of the production of karyotypically stable partial hybrids involving highly unrelated species from two subfamilies of the Gramineae (Pooideae — oat, and Panicoideae — maize) and the subsequent recovery of fertile oat-maize chromosome addition lines. These represent novel material for gene/ marker mapping, maize chromosome manipulation, the study of maize gene expression in oat, and the transfer of maize DNA, genes, or active transposons to oat.Joint contribution of the Minnesota Agricultural Experiment Station and USDA-ARS. Scientific journal series paper No. 21 859 of the Minnesota Agricultural Experiment Station. Mention of a trademark or proprietary product does not constitute a guarantee or warranty by the USDA-ARS or the University of Minnesota and does not imply approval over other products that also may be suitable  相似文献   

17.
A plant with 2n = 14 + 1 ring chromosomes was obtained in the progeny of a primary trisomie for chromosome 7 of a two-rowed cultivar, Shin Ebisu 16. The morphological characteristics of the trisomic plants with an extra ring chromosome were similar to the primary trisomic for chromosome 7 (Semierect), which suggests that it originated from this chromosome. The ring chromosomes were not completely stable in mitotic cells because of abnormal behavior. Chromosome complements varied in different plants and in different roots within a plant. Root tip cells and spikes with 2n = 14 and 14 + 2 ring chromosomes were observed on plants with 14 + 1 ring chromosomes. Breakage-fusion-bridge cycle was inferred. The ring chromosome was associated with two normal homologues forming a trivalent in 17.6% sporocytes at metaphase I. The transmission of the extra ring chromosome was 23.1% in the progeny of the plant with 14 + 1 ring chromosomes. Trivalent formation may have been much higher at early prophase stages which were difficult to analyze in barley; only 4 of 120 sporocytes analyzed showed an isolated ring at pachytene. The ring chromosome moved to one pole without separation in 24.7% of the sporocytes at AI, and divided in 27.1% sporocytes giving rise to 8-8 separation. Only 10% of the sporocytes showed bridge formation at AI.  相似文献   

18.
Z X Wang  N Iwata 《Génome》1995,38(4):696-705
Eight types of aneuhaploids (Aneuhaplo 4, 5, 6, 8, 9, 10, 11, and 12) and eight types of tetrasomics (Tetraplo 4, 5, 6, 7, 8, 9, 10, and 12) of rice have been obtained from anther culture of trisomics. This paper reports the plant morphology of these aneuploids and their chromosome behavior at metaphase I. Aneuhaploids for different chromosomes are distinguishable from each other and are morphologically similar to the parental trisomics, suggesting that the extra chromosome has similar genetic effects on plant morphology at the haploid level as at the diploid level. Similarly, tetrasomics with different extra chromosomes are distinguishable from each other and are similar morphologically to the parental trisomic. However, stronger changes in morphological characters were observed in tetrasomics compared with trisomics having the same extra chromosome, as a result of a dosage effect of the extra chromosomes. Comparing plant size between aneuhaploid, tetrasomic, and trisomic with the same extra chromosome, it was shown that the trisomic was the largest, the tetrasomic was of medium size, and the aneuhaploid was the smallest, except for those plants with an extra chromosome 8 in which plant size is dramatically decreased in both the aneuhaploid and the tetrasomic. At metaphase I, aneuhaploids showed chromosome configurations of 1 II + 11 I and 13 I. The frequency of the 1 II + 11 I configuration is higher than 70%, indicating that homologous chromosomes in aneuhaploids tend to stay associated in meiosis. Intragenome chromosome pairing (2 II + 9 I), so called secondary association, was observed in the aneuhaploid for chromosome 5. Tetrasomic plants showed 5 kinds of chromosome configurations: 1 IV + 11 II, 1 III + 11 II + 1 I, 13 II, 12 II + 2 I, and 11 II + 4 I. A chromosome configuration of 13 II was often observed in tetrasomics with shorter extra chromosomes and a chromosome configuration of 1 IV + 11 II was often observed in tetrasomics with longer extra chromosomes. Aneuhaploids had complete seed sterility. Tetrasomics showed very poor pollen fertility and complete seed sterility, except for a few shriveled seeds that were observed in Tetraplo 6 and 9. This is the first report in rice where many aneuhaploids and tetrasomics have been characterized. This information will help to further unravel rice aneuploidy and cytogenetics. The aneuploids obtained here will be very useful tools for the study of genetics and breeding in rice.  相似文献   

19.
Four tertiary trisomic plants are reported here, two of them (Nos. Tr11 and Tr13) from selfed progeny of a triploid Pearl millet and the other two (Nos. 3/12 and 16/7) from the progenies of radiation induced interchange heterozygotes. The extra chromosome in Tr13 and 3/12 was the nucleolus organizing chromosome. In No. 16/7 an extra chromosome enters into an association chromosomes were also involved. Meiotic behaviour in these four trisomics indicates that Tr11 and 3/12 are tertiary trisomics. It is suggested that two reciprocal translocations have occurred between two sets of chromosomes in the triploid parent and that syngamy has taken place in such a way that four interchange chromosomes and one non-interchange nucleolus organizing chromosome have come together in the offspring. The extra chromosome in No. 16/7 is an interchange chromosome which is homologous to one of the chromosomes of an interchange complex of six chromosomes.  相似文献   

20.
R J Singh  T Tsuchiya 《Génome》1993,36(2):350-355
The origin, identification, meiotic chromosome behavior, and breeding behavior of an unstable trisomic barley were studied. The extra chromosome originated by breakage and fusion of an acrocentric chromosome 3 in a plant from an F2 population of a cross between acrotrisomic 3L3S (2n = 14 + 1 acro3L3S) and a balanced lethal stock, xc. (xantha) ac (albino). The F2 population segregated only for the albino trait. The genotypic constitution of the trisomic plant was ac ac (for both normal chromosome 3) and Ac (for the unstable metacentric chromosome). The unstable extra metacentric chromosome was designated as metacentric 3B (abbreviated as meta3B). Meiotic chromosome behavior in plants with 2n = 14 + 1 meta3B differed from plant to plant and within spikes. Some plants showed only trisomic cells with a chromosome configuration of 1 III + 6 II and 7 II + 1 I at metaphase I, whereas other plants showed both trisomie and disomic cells (7 II) that resulted from the elimination of the extra meta3B. The frequency of ring trivalents was low (6.8%). An average transmission rate of unstable meta3B ranged from 4.3 to 12.9%. The elimination of meta3B, and hence loss of the dominant Ac allele, resulted in albino seedlings as well as white stripes on plants, leaves, and spikes. Chromosome numbers of albino seedlings in the progeny of 2n = 14 + 1 meta3B were all diploid (2n = 14), while green seedlings contained 2n = 14 + 1 meta3B. However, progenies of some spikes of one trisomic plant showed a low frequency of green diploids and metatrisomics (2n = 14 + 1 meta3B), which was attributed to crossing-over.  相似文献   

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