Pistillate-parental differences and the ability to produce interspecific hybrids between Lycopersicon esculentum and 'peruvianum-complex' species (L. peruvianum and L. chilense)

To overcome the cross-breeding barriers between the cultivated tomato Lycopersicon esculentum and the ‘peruvianum-complex’, hybrid production ability (HPA) and pistillate-parental differences were investigated. As a criterion of HPA, the number of germinated ovules per fruit (GPF) was used. GPF was expressed as GPF = OPF × GPO, where OPF is the number of ovules per fruit, and GPO is the proportion of germinated ovules to total ovules obtained. The interspecific crossing between nine varieties and three ‘peruvianum-complex’ accessions revealed that the cultivars ‘Sekaiichi’,‘Ponde Rooza’ and ‘Early Pink’ showed quite high and stable GPF over the years, but the cv.‘Best of All’ produced no hybrids. Variance analysis for GPF, OPF and GPO, and their correlation with seven sexual organ morphological traits and three fruit morphological traits were performed. These results indicated that choosing both the pistillate parents with wider reproductive organs for high OPF and appropriate environmental conditions for high GPO might be significant for enhancing GPF in interspecific crossing.     

Overcoming unilateral breeding barriers between Lycopersicon peruvianum and cultivated tomato,Lycopersicon esculentum

Summary Treatment of Lycopersicon peruvianum stigmas with an artificial medium analogous to stigmatic exudate allowed pollen germination and growth on immature pistils. Growth of Lycopersicon esculentum pollen tubes to L. peruvianum ovules, an otherwise incompatible cross, was achieved following such treated bud-pollinations. No plantlets were recovered, although a few embryos from this cross at the globularity heart stage of development were excised at 22 days after pollination, indicating the presence of crossing failures as severe as in the reciprocal cross. Hybrid plants were obtained from the reciprocal cross, using as pollen parent an L. peruvianum line selected for congruity with L. esculentum. Bud pollinations to L. peruvianum, using these interspecific F₁ hybrids as the pollen parent, allowed viable embryo development and plantlet recovery. Resulting backcross plants, which possess approximately 1/4 L. esculentum genome in a L. peruvianum cytoplasm, may facilitate further introgression of the L. esculentum nuclear genes into in foreign cytoplasm.     

Transfer of powdery mildew resistance from Brassica carinata to Brassica oleracea through embryo rescue

Interspecific hybrid plants and backcross 1 (BC₁) progeny were produced through sexual crosses and embryo rescue between Brassica carinata accession PI 360883 and B. oleracea cvs Titleist’and‘Cecile’to transfer resistance to powdery mildew to B. oleracea. Four interspecific hybrids were obtained through application of embryo rescue from crosses with B. carinata as the maternal parent, and their interspecific nature confirmed through plant morphology and random amplified polymorphic DNA (RAPD) analysis. Twenty‐one BC₁ plants were obtained through sexual crosses and embryo rescue although embryo rescue was not necessary to produce first backcross generation plants between interspecific hybrids and B. oleracea. All interspecific hybrids and eight of the BC₁ plants were resistant to powdery mildew.     

Variation for aphid resistance and insecticidal acyl sugar expression among and within Lycopersicon pennellii-derived inbred backcross lines of tomato and their F1 progeny

Five inbred backcross lines (IBL) were selected for higher relative expression of insecticidal acyl sugars (rank average) from an inbred backcross population derived from the cross Lycopersicon esculentum cultivar ‘Peto 84’×Lycopersicon pennellii accession LA716. These five BC₂S₅ IBLs were crossed in a partial diallel design (Method II), and their self and F₁ progeny and three control cultivars were tested at two California field locations in 1996. Counts of potato aphids, Macrosiphum euphorbiae Thomas, on leaves, as a measure of plant infestation, revealed significant general combining ability (GCA) for lower aphid numbers with IBL44 and IBL59; the F₁ hybrid IBL44 × IBL59 had significantly fewer aphids per leaflet than the susceptible cultivar ‘Alta’. GCA for acyl sugars was associated with IBL59 only. Of all the IBL and IBL × IBL F₁ hybrids, only IBL59 produced significant levels of acyl sugars. Significant within IBL59 variation for acyl sugars was observed, but not for aphid resistance. Our results suggest that factors other than acyl sugars contributed to L. pennellii-derived aphid resistance in IBL × IBL F₁ hybrids and IBL that do not produce significant amounts of acyl sugars. IBL59 and IBL44 may be useful for breeding for aphid resistance in cultivated tomato.     

Production of interspecific F1 hybrids, BC1, BC2 and BC3 populations between Lycopersicon esculentum and two accessions of Lycopersicon peruvianum carrying new root-knot nematode resistance genes

Two accessions of Lycopersicon peruvianum Mill. (PI270435, PI126443) carrying Mi-2 and Mi-3, respectively, new root-knot nematode resistance genes, were selected as the male parents for crosses with L. esculentum Mill. in order to produce interspecific hybrids. Crossability barriers between these two distantly related species were circumvented by ovule culture. A total of ten interspecific F₁ hybrid plants were produced. The hybrid nature of the putative F₁ plants was confirmed by a comparison of several morphological characteristics and a PCR-based assay. Eight of ten hybrid plants were backcrossed with L. esculentum to generate a total of 98 BC₁ progeny. Two lines were advanced to the BC₂ and BC₃ levels. Based on these results, ovule culture was found to be an effective method for the production of novel interspecific F₁ and BC₁ plants. This revised version was published online in July 2006 with corrections to the Cover Date.     

Combining ability for beet armyworm, Spodoptera exigua, resistance and horticultural traits of selected Lycopersicon pennellii-derived inbred backcross lines of tomato

Ten selected inbred backcross lines (IBL), from a Lycopersicon esculentum cv.‘Peto 84’×Lycopersicon pennellii IBL population, with resistance to beet armyworm (BAW), Spodoptera exigua, higher fruit mass and fruit yield, were crossed with eight elite cultivated L. esculentum inbred lines in a Design II mating design. Three elite inbreds were also crossed to ‘Peto 84′, the IBL recurrent parent, as a control for combining ability. Field plots of all resulting F₁ progenies and control cultivars were inoculated with BAW eggs and evaluated for resistance to BAW, fruit mass, fruit yield, vine size and maturity at three field locations. Reductions in fruit damage by BAW were found in four of the 10 IBL F₁ progenies. Significant male and female general combining ability (GCA) estimates for BAW resistance were observed, but significant specific combining ability for BAW resistance was not detected. The fruit mass of F₁ hybrids was significantly lower than large-fruited controls, but was not significantly different from elite inbred by ‘Peto 95’F₁ hybrids. Selection based on inbred performance identified IBL with positive GCA for BAW resistance and yields in Design II hybrids. BAW resistance in the 09 selected IBL and IBL- derived F₁ progeny was associated with two undesirable traits, later maturity and larger vine size. Index selection of IBL was more effective at identifying IBL with positive GCA for fruit mass and fruit yield than GCA for BAW resistance.     

Interactions of BA, GA3, NAA, and surfactant on interspecific hybridization of Lycopersicon esculentum × L. chilense

Time consuming and expensive tissue culture type techniques are currently used for achieving Lycopersicon esculentum × L. chilense interspecific hybrids. The objective of this study was to determine if the number of viable hybrid seed produced directly from this wide cross could be improved by optimizing female/male parent selection when treated with various hormones in the presence or absence of a surfactant (Triton X-100). Individual or combination treatments of BA(6-benzylaminopurine), GA₃ (gibberellic acid), and/or NAA(α-naphthaleneacetic acid) were applied (both with and without surfactant)to the ovaries of two L. esculentum female parents (Fla 7613 and89S) one day following pollination by nineL. chilense accessions. Hormone applications over 1-, 3-, and 5-dayperiods were made and all treatments were compared to a distilled water control. Across all experiments 21 viable hybrids from 1920 fruits (1.1%) were obtained from2128 pollinations. There were five consistent trends observed across three experiments. First, treatments utilizing NAA produced more hybrids than any other treatment hormone. Second, eight of the total 21 hybrids resulted from five consecutive days of hormone treatments (P-value ≤0.01), and six of those eight were realized when NAA was involved. Third, the use of Triton X-100 in combination with the hormone treatments significantly reduced the number of fruits/pollination (P-value ≤ 0.01), but did not significantly reduce the number of overall viable hybrids (P-value > 0.4). Fourth, the L. chilense male parent LA2759 significantly produced the most interspecific hybrids (P-value ≤ 0.04),seven hybrids from 295 fruits, compared to the male parent LA130 which did not produce any hybrids (from 293 fruits) -other accessions were intermediate. Fifth, nearly ten times more hybrids were produced from crosses with Fla 7613 than with 89S. These experiments provide a foundation for future studies to discern specific concentrations, timing and frequency of application of NAA and possibly, GA₃, to efficiently produce interspecific hybrids of L. esculentum × L. chilense. This revised version was published online in August 2006 with corrections to the Cover Date.     

Interspecific hybrids between three eggplant (Solanum melongena L.) cultivars and two wild species (Solanum torvum Sw. and Solanum sisymbriifolium Lam.)

Three Greek eggplant cultivars, ‘Langada’, ‘Tsakoniki’ and ‘Emi’ (2n= 24), were crossed with two wild species (Solanum torvum Sw., 2n= 24 and Solanum sisymbriifolium Lam., 2n= 24). Ovules isolated 15-27 days after pollination were cultured in a modified MS medium at 24°C and a 16h photoperiod. Fifty days later, the ovules were dissected and the interspecific embryos were cultured in the same medium. Interspecific hybrids were achieved only from crosses between the eggplant cultivars and S. torvum. The hybridity of the putative interspecific F¹ hybrid (Solanum melongena×S. torvum) was confirmed by using morphological and biochemical (isozyme isocitrate dehydrogenase A, phosphoglucomutase A, phosphoglucose isomerase B, 6-phosphogluconate dehydrogenase A, 6-phosphogluconate dehydrogenase B) markers. The F¹ plants (‘Langada’×S. torvum) were selfpollinated and backcrossed to both parents. Fruits, however, were produced only when the F¹ hybrid was backcrossed as female with the eggplant cultivar ‘Langada’.     

Cytological and Isozyme Evaluation of Tall Fescue × Italian Ryegrass Hybrids

Development of tall fescue (Festuca arundinacea Schreb., 2n = 6x - 42) × Italian rye-grass (Lolium multiflorum Lam., 2n = 2 ×= 14) hybrids would enhance efforts to improve the quality of tall fescue. Two ‘Kentucky 31’ tall fescue בLemtal Italian ryegrass hybrids were obtained via embryo rescue on MS media containing casein hydrolysate, ascorbic acid and sucrose. Chromosome pairing at metaphase I had an average of more than 12 bivalents per cell. Since Festuca-Lolium pairing can account only for seven of the paired chromosomes, intergenomic as well as interspecific chromosome pairing is indicated. There was no cytoplasmic effect on chromosome pairing. To determine if enzymes could be used as genetic markers for distinguishing hybrids from self-contaminants in crosses, zymograms of PGI, 6-PGD, MDH, GOT and ACPH were obtained from parents and hybrids using starch gel etectrophoresis. PGI, 6-PGD and MDH had fewer bands in the diploid ryegrass, as compared with the hexaploid tall fescue and the tetraploid hybrid.     

Attempts to overcome postfertilization barrier in interspecific crosses of the genus Lens

Wild relatives are a potential source of genetic diversity to lentil (Lens culinaris Medik). The objective of this research was to obtain viable interspecific hybrids between the domesticated lentil and its wild relatives. The paper details the results of a number of interspecific crosses among L. culinaris, L. orientalis, L. odemensis, L. ervoides and L. nigricans. Viable hybrids were produced between L. culinaris × L. orientalis, L. culinaris × L. nigricans, L. culinaris × L. ervoides and between L. culinaris × L. odemensis. Further viable hybrids were obtained between L. culinaris and L. ervoides, which have the potential to be a ‘bridge’ in hybridization to L. culinaris for specific L. nicrigans lines which proved recalcitrant in L. culinaris × L. nigricans crosses. This is the first time that four wild species of lentils have been used successfully in hybridization with cultivated lentils, and viable hybrids produced. This paper also suggests that the artificial supplement of GA₃, hormone is needed after fertilization for the normal growth of the hybrid embryo, possibly as the natural GA₃ production is restricted with alien pollinations in cultivated lentils in both F₁ and backcross hybrids.     

Plant Regeneration from the Hybridization of Brassica juncea and B. napus Through Embryo Culture

Crosses were made to produce interspecific hybrids between Brassica napus × B. juncea and their reciprocals with the aid of embryo culture techniques. A better response of hybrid embryo culture was obtained from two cross combinations of B. juncea × B. napus (Ames 24521 × Huyou 15 and Vittasso × Zheshuang 72) than from their reciprocals. Embryo culture was more effective in terms of plant regeneration when embryos were cultured in vitro at 15 days after pollination (DAP), while more calli were initiated when embryos were excised and cultured at 10 DAP. A better response was observed on the MS medium with 0.3 mg l^?1 naphthylacetic acid (NAA) + 1.5 mg l^?1 6‐benzylaminopurine (BAP) and with 0.3 mg l^?1 NAA + 2.0 mg l^?1 BAP. Callus formation and plant regeneration on these two media reached 55.43 and 26.65 %, and 66.98 and 24.61 %, respectively.     

Zingiberene-mediated resistance to the South American tomato pinworm derived from Lycopersicon hirsutum var. hirsutum

The Lycopersicon hirsutum var. hirsutum accession PI 127826 is recognized as a good source of resistance to arthropod pests due to the action of the allelochemical zimgiberene, a sesquiterpene present in its glandular trichomes. Five genotypes were selected from the F₂ generation of the interspecific cross Lycopersicon esculentum ‘TOM-556’ × Lycopersicon hirsutum var. hirsutum ‘PI 127826’, based on their low levels (BPX-368-clone#56) or high levels(BPX-368-clone#92, BPX-368-clone#105,BPX-368-clone#179, BPX-368-clone#250) of zingiberene. The five F₂ genotypes were tested for resistance to the South American tomato pinworm Tuta absolutaalong with accession L. esculentum ‘TOM-556’ (pinworm susceptible), and the accessions L. hirsutum var. hirsutum ‘PI 127826’ and L. pennellii ‘LA716’ (resistant). The F₂ clones selected for high foliar zingiberene levels showed lower scores for leaflet lesion type(LLT), percent leaflets attacked (PLA) and overall plant damage (OPD) than the low zingiberene genotypes. The results indicated that zingiberene mediates resistance to the South American pinworm, based on feeding and on ovipositing deterrence, in populations derived from the interspecific cross between Lycopersicon. esculentum and Lycopersicon hirsutum var. hirsutum. Indirect selection for high foliar zingiberene content is suggested as an efficient technique for breeding tomatoes for resistance to the South American tomato pinworm. This revised version was published online in July 2006 with corrections to the Cover Date.     

Interspecific hybridization in sexual diploid Allium senescens var. minor× apomictic tetraploid A. nutans and sexual diploid A. senescens var. minor× apomictic hexaploid A. senescens

Fourteen interspecific hybrids in sexual diploid Allium senescens var. minor× apomictic tetraploid Allium nutans L. crosses, and eight interspecific hybrids in sexual diploid A. senescens var. minor× apomictic hexaploid A. senescens L. crosses were produced. The number of chromosomes was 2n= 24 in interspecific hybrids of diploid × tetraploid, and 2n= 32 in diploid × hexaploid crosses. Triploid and tetraploid interspecific hybrids showed intermediate parental morphological characteristics. Tetraploid interspecific hybrids of A. senescens var. minor×A. senescens crosses formed two groups based on leaf colour and leaf width. Seeds were formed in 11 out of 14 triploid interspecific hybrids under natural conditions. In cytological observations of parthenogenesis, three out of 12 triploid interspecific hybrids and five out of eight interspecific tetraploid hybrids were observed. Parthenogenesis ranged from 26.0% to 86.0% in five tetraploid interspecific hybrids. Non-parthenogenesis to parthenogenesis segregated in a 3:5 ratio in A. senescens var. minor×A. senescens crosses.     

The use of ISSR markers to identify Texas bluegrass interspecific hybrids

Seventeen ISSR primers were screened on Texas bluegrass (Poa arachnifera), Kentucky bluegrass (P. pratensis), Canadian bluegrass (P. compressa), Argentine bluegrass (P. ligularis), cv. ‘Sherman’ (P. secunda), putative Texas × Kentucky (TK) (P. arachnifera ×P. pratensis) hybrids and hybrids involving Texas, Canadian, and Sherman bluegrass [(P. arachnifera × P. compressa) × (P. arachnifera ×P. secunda)], to determine whether they could be used to produce robust and reproducible DNA fingerprints and identify interspecific hybrids. Nine of the 17 primers consistently produced robust fingerprints and nine 2‐way primer combinations were also selected. DNA fingerprints were highly reproducible and the majority of the selected primers (16/18) amplified hybrid profiles using two putative TK full‐sib hybrids. Combined with a rapid DNA extraction protocol, the ISSR technique enabled a fast and practical way to detect F₁ interspecific hybrids early in the breeding programme and could also be useful for other applications that require DNA‐based markers.     

Trichomes on leaves of Lycopersicon hirsutum,L. esculentum and their hybrids

Summary Density of certain types of trichomes were characterized on leaflets of Lycopersicon hirsutum, L. esculentum and their hybrids. Trichome density covaried with leaflet development, and depending on the species and type of trichome, density also covaried with leaflet surface. Type IV trichomes were not observed on leaflets of L. esculentum and Type V trichomes were not observed on leaflets of L. hirsutum. Type IV and V trichomes were present on F₁ hybrids and densities of these two types of trichomes segregated in the F₂ population. When corrected for differences of leaflet size between L. hirsutum and L. esculentum, densities of Type VI trichomes were more similar between species than uncorrected densities. There was little difference in Type VI density among F₂ individuals. The appearance, lipid and phenol staining properties, and native fluorescence of the multicellular tip of the Type VI trichome differed between L. hirsutum and L. esculentum. F₁ hybrids were intermediate for these characters and these characters segregated in the F₂ generation.The investigation reported in the paper (84-10-94) is in connection with a project of the Kentucky Agricultural Experiment Station and is published with the approval of the Director. This material is based upon work supported, in part, by the U.S. Department of Agriculture, Agreement No. 59-2213-1-1-717-0 (Competitive Research Grants Program).     

Identification of barriers to interspecific crosses in the genus Trifolium

Summary A study of pre- and post-fertilisation barriers after interspecific crosses of diploid and tetraploid Trifolium pratense L. and wild species T. alpestre L., T. medium L. and T. sarosiense Hazsl. was aimed at finding of a promising cross combination for obtaining hybrids. The growth of pollen tubes was arrested in interspecific crosses mainly when T. pratense was at a diploid level. To investigate the post-fertilisation barriers in detail, the hybrid embryo viability was traced by two clearing treatments of immature seeds: (1) using chloral hydrate (which proved to be most appropriate); and (2) a mixture of benzyl benzoate and dibutly phthalate. In interspecific combinations T. pratense (4×) × either T. alpestre or T. sarosiense, enlargement of immature seeds occurred, but no hybrid embryo was traced. Of the wild species used as a male parent for crosses, T. medium was the only exception from the point of view of fertilisation. Globular, heart and the early torpedo stages of hybrid embryos were observed 7 days after pollination (DAP) but only when T. pratense was at a tetraploid level. When T. pratense (2×, 4×) was used as a male parent for interspecific crosses with T. alpestre, T. medium and T. sarosiense, strong defects in various stages of embryogenesis were observed, particularly wrinkled and narrowing embryo sacs caused by an expansion of endothelial cells. We conclude with the following finding: (1) to make crosses only in one direction with T. pratense as a female parent and T. medium as a male; (2) to use tetraploid plants of T. pratense; (3) and to excise hybrid embryos at an early torpedo stage, about 7 DAP.     

Genetics of Resistance to Fruit Borer, Heliothis armigera Hübner, in Lycopersicon

Inheritance of resistance to the fruit borer, Heliothis armigera Hübner in tomato was studied in TWO interspecific crosses. L. hirsutum f. gLabratum‘B 6013′. a fruit borer-resistant parent, was crossed with two susceptible L. esculentum varieties ‘HS 101’ and. ‘HS 102′. Six generations of two crosses were evaluated for percentage fruit: infestation (on a number basis) and the estimates of the gene effects were derived from the generation mean using a three parameter model A preponderance of additive gene effects was found for this trait. Heritability in the narrow sense was observed to be high indicating the fixable nature of the character.     

Hybridization, Through Culture of Embryos and Immature Seeds, of a Range of Tomato Cultivars with a Tomato Somatic Hybrid [Lycopersicon esculentum (+) L. peruvianum]: Emergence of a Possible New Marker Gene for Tomato Breeding

Two hexaploid somatic hybrids [Sh; L. esculentum (+) L. Peruvianum] accessions 6 and 18 were back-crossed with two diploid L. Esculentum cultivars ‘Moneymaker’ (mm) and ‘pusa Ruby’ (pr). Twenty-two plants of the bc2 generation were produced by backcrossing 7 bcl plants (mm x sh, 6, 18) with five tomato cultivars. Fourteen of the bc2 plants were self-fertile, five produced anther cones with anthocyanin pigmentation not present in the parents. A bc3 generation was developed by crossing the four cultivars as female parent with three bc2 generation plants. The bc3 progeny derived from one pollen parent plant were produced without the need to culture immature seeds. They segregated with respect to pigmented anther cones and were self-fertile. The anther cone pigmentation of the pollen parent plant was associated with increased seed set, greater fruit size and an orange-red fruit colour. These features were transmitted to the fertile bc3 generation. Conversely, bc3 offspring involving the other two parent plants were only recovered by culture of immature seeds. The recovery of diploid plants in BCl and self-fertility in BC2 resulted in almost total recovery of the tomato cultivar characteristics (fruit size, colour and number of seeds) by BC3.     

Characterization of a green non‐dormant sunflower (Helianthus annuus) mutant NDG

A sunflower (Helianthus annuus L.) mutant was observed in the progeny of a cross between the sunflower cultivar ‘HA 89’ and an amphiploid of a Helianthus divaricatus L. × P21 cross that exhibited loss of dormancy induction in the developing embryo. Seeds of this mutant frequently germinate on the head about 40 days after pollination. The cotyledons of this mutant remain green, whereas some other non‐dormant mutants exhibit loss of pigmentation. The objectives of this investigation were to compare levels and activities of abscisic acid (ABA), a plant hormone that induces dormancy in developing embryos, in the non‐dormant green mutant (NDG) and ‘HA 89’ from which NDG was derived. Immunoassays showed that abscisic acid was present in NDG and the levels were not significantly different from those in ‘HA 89’. Exposure of excised NDG mutant embryos to 40 μm abscisic acid failed to prevent germination, suggesting that non‐dormancy could result from impairment in ABA receptors or from a defect in other proteins participating in the subsequent signalling pathway that normally induces dormancy.     

Somatic hybridization as a tool for tomato breeding

Protoplast fusion can be used to produce somatic hybrids of species that cannot be obtained by sexual hybridization. The possibility to introgress genes from Solanum species into the cultivated tomato species Lycopersicon esculentum, and to obtain novel cytoplasm-nucleus combinations (cybrids) was considered as an important strategy to extend the genetic variation available for tomato breeding. Somatic hybrids between L. esculentum and other Lycopersicon species, as well as between L. esculentum and Solanum or Nicotiana species, have been produced. Specific mutants, genotypes with antibiotic resistances, and metabolic inhibition by iodoacetate or iodoacetamide and irradiation were used for the selection of hybrids. In addition, the improvement of protoplast culture techniques and the use of the favourable tissue culture traits derived from species such as L. peruvianum, which have been introduced into tomato by classical breeding, allowed the efficient recovery of somatic hybrids. However, the occurrence of somatic incongruity in fusion combinations of L. esculentum and Solanum and even more in L. esculentum and Nicotiana, did not allow the production of true cybrids and/or fertile hybrids, indicating the importance of both cytoplasm-nucleus and nucleus-nucleus interactions in somatic incongruity. Another problem with fusions between distantly related species is the strongly reduced fertility of the hybrids and the very limited homoeologous recombination between chromosomes of the parental species. Partial genome transfer from donor to recipient through microprotoplast (+) protoplast fusion, and the production of monosomic or disomic chromosome addition lines, light overcome some of these problems. In symmetric somatic hybrids between L. esculentum and S. tuberosum the occurrence of limited somatic and meiotic recombination was demonstrated. Fertile progeny plants could be obtained, though at a low frequency, when embryo rescue was performed on a large scale after backcrossing hexaploid somatic tomato (+) potato hybrids with a tetraploid potato genotype. The potential value of genomic in situ hybridization (GISH) and RFLPs for the analysis of the genome/chromosome composition of the hybrids has been demonstrated for intergeneric somatic hybrids between Lycopersicon and Solanum.Abbreviations cpDNA chloroplast DNA - mtDNA mitochondrial DNA