WO2018159096A1 - Strawberry having fragaria iinumae-derived cytoplasm and showing gene-cytoplasmic male sterility - Google Patents
Strawberry having fragaria iinumae-derived cytoplasm and showing gene-cytoplasmic male sterility Download PDFInfo
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- WO2018159096A1 WO2018159096A1 PCT/JP2017/047418 JP2017047418W WO2018159096A1 WO 2018159096 A1 WO2018159096 A1 WO 2018159096A1 JP 2017047418 W JP2017047418 W JP 2017047418W WO 2018159096 A1 WO2018159096 A1 WO 2018159096A1
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- strawberry
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- fragalia
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- 241000220223 Fragaria Species 0.000 title claims abstract description 101
- 235000011363 Fragaria x ananassa Nutrition 0.000 title claims abstract description 92
- 235000016623 Fragaria vesca Nutrition 0.000 title claims abstract description 90
- 206010021929 Infertility male Diseases 0.000 title claims abstract description 46
- 208000007466 Male Infertility Diseases 0.000 title claims abstract description 46
- 210000000805 cytoplasm Anatomy 0.000 title claims abstract description 18
- 240000001409 Fragaria iinumae Species 0.000 title claims abstract description 7
- 235000016952 Fragaria iinumae Nutrition 0.000 title claims abstract description 4
- 238000004519 manufacturing process Methods 0.000 claims abstract description 12
- 108090000623 proteins and genes Proteins 0.000 claims description 37
- 230000001086 cytosolic effect Effects 0.000 claims description 33
- 241000894007 species Species 0.000 claims description 23
- 238000010899 nucleation Methods 0.000 claims description 8
- 241000196324 Embryophyta Species 0.000 claims description 7
- 238000000034 method Methods 0.000 claims description 7
- 210000000349 chromosome Anatomy 0.000 claims description 5
- 240000009088 Fragaria x ananassa Species 0.000 claims description 2
- 235000021012 strawberries Nutrition 0.000 description 11
- 230000035558 fertility Effects 0.000 description 9
- 230000002159 abnormal effect Effects 0.000 description 8
- 238000009395 breeding Methods 0.000 description 5
- 230000001488 breeding effect Effects 0.000 description 5
- 230000008722 morphological abnormality Effects 0.000 description 5
- IAKHMKGGTNLKSZ-INIZCTEOSA-N (S)-colchicine Chemical compound C1([C@@H](NC(C)=O)CC2)=CC(=O)C(OC)=CC=C1C1=C2C=C(OC)C(OC)=C1OC IAKHMKGGTNLKSZ-INIZCTEOSA-N 0.000 description 4
- 230000005856 abnormality Effects 0.000 description 4
- 230000013011 mating Effects 0.000 description 4
- 238000011084 recovery Methods 0.000 description 4
- 235000013399 edible fruits Nutrition 0.000 description 3
- 230000002438 mitochondrial effect Effects 0.000 description 3
- 210000000056 organ Anatomy 0.000 description 3
- 208000020584 Polyploidy Diseases 0.000 description 2
- 230000015572 biosynthetic process Effects 0.000 description 2
- 229960001338 colchicine Drugs 0.000 description 2
- 239000000463 material Substances 0.000 description 2
- 210000004940 nucleus Anatomy 0.000 description 2
- 210000001519 tissue Anatomy 0.000 description 2
- 241000908846 Fragaria nilgerrensis Species 0.000 description 1
- 241000238631 Hexapoda Species 0.000 description 1
- 108700005079 Recessive Genes Proteins 0.000 description 1
- 102000052708 Recessive Genes Human genes 0.000 description 1
- 210000004027 cell Anatomy 0.000 description 1
- 239000003086 colorant Substances 0.000 description 1
- 238000011109 contamination Methods 0.000 description 1
- 201000010099 disease Diseases 0.000 description 1
- 208000037265 diseases, disorders, signs and symptoms Diseases 0.000 description 1
- 238000010413 gardening Methods 0.000 description 1
- 239000003550 marker Substances 0.000 description 1
- 230000008774 maternal effect Effects 0.000 description 1
- 230000000877 morphologic effect Effects 0.000 description 1
- 238000010606 normalization Methods 0.000 description 1
- 230000008520 organization Effects 0.000 description 1
- 230000001954 sterilising effect Effects 0.000 description 1
- 238000004659 sterilization and disinfection Methods 0.000 description 1
Images
Classifications
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- A—HUMAN NECESSITIES
- A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
- A01H—NEW PLANTS OR NON-TRANSGENIC PROCESSES FOR OBTAINING THEM; PLANT REPRODUCTION BY TISSUE CULTURE TECHNIQUES
- A01H1/00—Processes for modifying genotypes ; Plants characterised by associated natural traits
- A01H1/02—Methods or apparatus for hybridisation; Artificial pollination ; Fertility
- A01H1/021—Methods of breeding using interspecific crosses, i.e. interspecies crosses
-
- A—HUMAN NECESSITIES
- A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
- A01H—NEW PLANTS OR NON-TRANSGENIC PROCESSES FOR OBTAINING THEM; PLANT REPRODUCTION BY TISSUE CULTURE TECHNIQUES
- A01H1/00—Processes for modifying genotypes ; Plants characterised by associated natural traits
- A01H1/02—Methods or apparatus for hybridisation; Artificial pollination ; Fertility
- A01H1/022—Genic fertility modification, e.g. apomixis
- A01H1/023—Male sterility
-
- A—HUMAN NECESSITIES
- A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
- A01H—NEW PLANTS OR NON-TRANSGENIC PROCESSES FOR OBTAINING THEM; PLANT REPRODUCTION BY TISSUE CULTURE TECHNIQUES
- A01H1/00—Processes for modifying genotypes ; Plants characterised by associated natural traits
- A01H1/06—Processes for producing mutations, e.g. treatment with chemicals or with radiation
- A01H1/08—Methods for producing changes in chromosome number
-
- A—HUMAN NECESSITIES
- A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
- A01H—NEW PLANTS OR NON-TRANSGENIC PROCESSES FOR OBTAINING THEM; PLANT REPRODUCTION BY TISSUE CULTURE TECHNIQUES
- A01H6/00—Angiosperms, i.e. flowering plants, characterised by their botanic taxonomy
- A01H6/74—Rosaceae, e.g. strawberry, apple, almonds, pear, rose, blackberries or raspberries
Definitions
- the present invention relates to, for example, a strawberry having a nuclear gene cytoplasmic male sterility and a method for producing the strawberry.
- F1 varieties hybrid first breeding (hybrid first breeding (F1) varieties)
- “Yotsuboshi” jointly nurtured by the Agricultural Research Organization
- F1 varieties are expected to produce a new strawberry industry, such as production and sales of seeds and strawberry seedlings, in addition to high productivity due to hybrid strength, high growth rate and easy disease-free seedling production.
- male sterilization and mating are carried out manually at the time of F1 seeding, increasing the price of seeds and lowering the purity of seeds due to omission of males are problematic. Therefore, practical use of a strawberry mating mother having male sterility is desired in order to increase the efficiency of F1 seeding with high purity.
- Strawberries grown in runners can be used as ornamental / ground covers, and both white flowers and red flowers are commercially available, but both are single. Thus, conventionally, there is no double-flowering variety among ornamental strawberry varieties. In addition, when the existing variety is used for ground cover applications, it is necessary to remove the decayed fruits over time.
- an object of the present invention is to provide a strawberry mating mother having male sterility in order to improve the efficiency of F1 seeding with high purity.
- a strawberry having a combination of the cytoplasm derived from Fragalia iinumae and the nuclear genome of a strawberry cultivar becomes a nuclear gene cytoplasmic male sterile and completed the present invention. It came to do. That is, the present invention includes the following. (1) A strawberry or progeny having cytoplasm derived from Fragaria iinumae and having a nuclear gene cytoplasmic male sterility.
- the strawberry or its progeny according to (1) which has a nuclear genome of a strawberry cultivated species.
- FIG. 1 It is a figure which shows the production method of the strawberry strain
- the stamen morphological abnormality (petaloid male sterility) individual which appeared by continuously backcrossing a strawberry cultivar to a 10-ploid interspecific hybrid 08IK5804-01 having female fertility and pollen forming ability is shown. It shows stamen morphological abnormalities appearing in backcross individuals with cytoplasm derived from fragalia iinumae.
- the strawberry strains of various organ shapes and flower colors produced from stamen morphological abnormalities (petaloid type male sterility) lines having cytoplasm derived from fragalia iinumae are shown.
- Strawberries or progenies having the nuclear gene cytoplasmic male sterility (or cytoplasmic male sterility) according to the present invention have a cytoplasm derived from a diploid wild species, Fragalia iinumae.
- the nuclear gene cytoplasmic male sterility is a maternally inherited trait caused by a unique orf present on the mitochondrial genome. The trait is counteracted by the fertility restoration gene present in the nuclear genome, and pollen is normally formed.
- a fertility-recovering gene in the nuclear genome of Fragalia iinumae but a strawberry having a combination of cytoplasm derived from Fragalia iinumae and the nuclear genome of a strawberry cultivar is said to be a nuclear gene cytoplasmic male sterile. Based on knowledge.
- Strawberry cultivars from which the nuclear genome possessed by the strawberry having nuclear gene cytoplasmic male sterility according to the present invention are derived for example, Fragalia x Ananassa (octaploid cultivar), Fragalia x Ananassa and diploid wild species Fragalia nilgrensis And interspecific hybrids (deploid cultivated species; for example, cultivar “Momochi”) and the like.
- the progeny of a strawberry having a nuclear gene cytoplasmic male sterility according to the present invention refers to a strawberry having a nuclear gene cytoplasmic male sterility according to the present invention as a seed parent and a strawberry variety that can be crossed with the strawberry as a pollen parent.
- strawberry varieties that can be crossed with strawberry having nuclear gene cytoplasmic male sterility according to the present invention include, for example, 'Sachinoka', 'Cullenberry', 'Kaorino', 'Ookimi', 'Momochi' , “Kurume IH4”, “Saikuhime” and the like.
- organs or tissues such as seeds, flowers, leaves, stems, roots, or these organs or tissues Cell or cytoplasm.
- FIG. 1 A method for producing a strawberry having a nuclear gene cytoplasmic male sterility according to the present invention is shown in FIG.
- i represents the mitochondrial genome of Fraglia iinumae
- I represents the chromosome of the nucleus of Fraglia iinumae
- a represents the mitochondrial genome of F. x ananassa
- A indicates the chromosome of the nucleus of Fragalia x Ananassa.
- a strawberry cultivated species is crossed using fragalia iinumae (diploid) as a seed parent to produce an interspecific hybrid having cytoplasm derived from fragalia iinumae.
- fragalia iinumae diploid
- strawberries with nuclear gene cytoplasmic male sterility that do not form mature pollen appear.
- individuals that do not form mature pollen can be selected as strawberries having the nuclear gene cytoplasmic male sterility according to the present invention.
- the strawberry having a nuclear gene cytoplasmic male sterility according to the present invention or its progeny is used as a seed parent, and the strawberry cultivated species that can be crossed with the strawberry or its progeny is crossed as a pollen parent, from the seed parent after the crossing F1 seeds can be collected.
- the progeny can be obtained by continuously backcrossing a strawberry variety having a useful trait (for example, fruit quality or seasonality) to a strawberry having a nuclear gene cytoplasmic male sterility according to the present invention or its progeny.
- An excellent line having useful traits and having a nuclear gene cytoplasmic male sterility can be obtained. Examples of strawberry varieties having such useful traits include “Mie Motomoto No. 1” and “A8S4-147”.
- the excellent line having the nuclear gene cytoplasmic male sterility can be used as a seed parent for obtaining F1 seeds.
- the strawberry having the nuclear gene cytoplasmic male sterility according to the present invention described above has the following characteristics.
- F1 seeding of strawberries according to the prior art is manually sterilized and mated, which is associated with high labor costs and the risk of contamination of self-grown seeds.
- a strawberry line having a nuclear gene cytoplasmic male sterility in which no pollen formation according to the present invention is used as a seed parent it is possible to reduce the cost and reliability by simply mixing the pollen parent line and releasing pollen-borne insects. F1 seeds can be obtained.
- a strawberry having a nuclear gene cytoplasmic male sterility according to the present invention is a petalized stamen and has a petaloid cytoplasmic male sterility in which mature pollen is not formed.
- stamen morphological abnormal line When hybrids of 10-ploid hybrids with the nuclear genome of iinumae were crossed, the appearance rate of stamen abnormalities was 1.5%, and normalization of stamens was observed in most seedlings (Table). 2).
- strawberry cultivars are crossed to stamen morphologically abnormal lines, many stamen morphologies are abnormal. When interspecific hybrids having the nuclear genome of iinumae are crossed, most of them are normalized. There is a recovery gene in the nuclear genome of iinumae, and it is considered possible to produce seedlings whose pollen-forming ability has been restored (recovery of fertility). Since stamens petalize in stamen morphologically abnormal lines, there are lines with white petals and double blooms.
- strawberry stamens with nuclear gene cytoplasmic male sterility according to the present invention have male sterility (petaloid male sterility) strawberry with morphological abnormalities (petalization) of different degrees and stamen petals. Can be used to produce double-flowered strawberries with high appreciation. All publications, patents and patent applications cited herein are hereby incorporated by reference in their entirety.
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Abstract
The purpose of the present invention is to provide a strawberry parent for crossing, said parent showing male sterility, for the efficient seed production of F1 with high purity. More specifically, the present invention relates to strawberry that has Fragaria iinumae-derived cytoplasm and shows gene-cytoplasmic male sterility.
Description
本発明は、例えば核遺伝子細胞質雄性不稔を有するイチゴ及びその作出方法に関する。
The present invention relates to, for example, a strawberry having a nuclear gene cytoplasmic male sterility and a method for producing the strawberry.
近年、農研機構共同育成の「よつぼし」を始め、種子繁殖型イチゴ品種（雑種第一代交配（Ｆ１）品種）育成が活発化している。Ｆ１品種は、雑種強勢による高生産性に加え、高い増殖率と容易な無病苗生産による大幅な栽培労力削減が期待され、種子やイチゴ苗の生産販売等、新たなイチゴ産業の創出につながる。しかしながら、Ｆ１採種時には除雄や交配を手作業で実施するため、種子の高価格化や、除雄漏れによる種子純度低下が課題となっている。
そこで、純度の高いＦ１採種の効率化のために、雄性不稔を有するイチゴ交配母本の実用化が望まれている。また、家庭園芸用として観賞用の赤花イチゴが販売されているが、より鑑賞価値の高いイチゴに対する要望も大きい。
通常のイチゴ品種は両性株であり、雌株品種が育成されたことがあるが、採種に利用されたことはない。唯一報告されている雄性不稔系統「ＭＳ−２」（核遺伝子型細胞質雄性不稔）（非特許文献１~３）の雄性不稔性は単因子劣性遺伝子支配であり、ホモ化すると後代がとれないため、育成中はヘテロ（表現形は可稔）個体を選抜する必要があり、選抜マーカーの開発を目指しているが、実用化には至っていない。
また、ランナーで増殖するイチゴは、観賞用／グランドカバーとして利用可能であり、白花、赤花ともに市販されているが何れも一重である。このように、従来において、観賞用イチゴ品種に八重咲き品種は存在しない。さらに、既存品種をグランドカバー用途に用いると結実するため、腐敗果を経時的に除去する必要がある。 In recent years, seed breeding type strawberry varieties (hybrid first breeding (F1) varieties), such as “Yotsuboshi” jointly nurtured by the Agricultural Research Organization, have been actively cultivated. F1 varieties are expected to produce a new strawberry industry, such as production and sales of seeds and strawberry seedlings, in addition to high productivity due to hybrid strength, high growth rate and easy disease-free seedling production. However, since male sterilization and mating are carried out manually at the time of F1 seeding, increasing the price of seeds and lowering the purity of seeds due to omission of males are problematic.
Therefore, practical use of a strawberry mating mother having male sterility is desired in order to increase the efficiency of F1 seeding with high purity. In addition, ornamental red strawberry is sold for home gardening, but there is a great demand for strawberry with higher appreciation value.
Normal strawberry varieties are amphoteric and female varieties have been bred but never used for seeding. The male sterility of the male sterility line "MS-2" (nuclear genotype cytoplasmic male sterility) (Non-patentDocuments 1 to 3), which is reported only, is dominated by a single factor recessive gene. Since it cannot be taken, it is necessary to select heterozygous individuals (with a phenotypic expression) during breeding, and aims to develop a selection marker, but has not yet been put into practical use.
Strawberries grown in runners can be used as ornamental / ground covers, and both white flowers and red flowers are commercially available, but both are single. Thus, conventionally, there is no double-flowering variety among ornamental strawberry varieties. In addition, when the existing variety is used for ground cover applications, it is necessary to remove the decayed fruits over time.
そこで、純度の高いＦ１採種の効率化のために、雄性不稔を有するイチゴ交配母本の実用化が望まれている。また、家庭園芸用として観賞用の赤花イチゴが販売されているが、より鑑賞価値の高いイチゴに対する要望も大きい。
通常のイチゴ品種は両性株であり、雌株品種が育成されたことがあるが、採種に利用されたことはない。唯一報告されている雄性不稔系統「ＭＳ−２」（核遺伝子型細胞質雄性不稔）（非特許文献１~３）の雄性不稔性は単因子劣性遺伝子支配であり、ホモ化すると後代がとれないため、育成中はヘテロ（表現形は可稔）個体を選抜する必要があり、選抜マーカーの開発を目指しているが、実用化には至っていない。
また、ランナーで増殖するイチゴは、観賞用／グランドカバーとして利用可能であり、白花、赤花ともに市販されているが何れも一重である。このように、従来において、観賞用イチゴ品種に八重咲き品種は存在しない。さらに、既存品種をグランドカバー用途に用いると結実するため、腐敗果を経時的に除去する必要がある。 In recent years, seed breeding type strawberry varieties (hybrid first breeding (F1) varieties), such as “Yotsuboshi” jointly nurtured by the Agricultural Research Organization, have been actively cultivated. F1 varieties are expected to produce a new strawberry industry, such as production and sales of seeds and strawberry seedlings, in addition to high productivity due to hybrid strength, high growth rate and easy disease-free seedling production. However, since male sterilization and mating are carried out manually at the time of F1 seeding, increasing the price of seeds and lowering the purity of seeds due to omission of males are problematic.
Therefore, practical use of a strawberry mating mother having male sterility is desired in order to increase the efficiency of F1 seeding with high purity. In addition, ornamental red strawberry is sold for home gardening, but there is a great demand for strawberry with higher appreciation value.
Normal strawberry varieties are amphoteric and female varieties have been bred but never used for seeding. The male sterility of the male sterility line "MS-2" (nuclear genotype cytoplasmic male sterility) (Non-patent
Strawberries grown in runners can be used as ornamental / ground covers, and both white flowers and red flowers are commercially available, but both are single. Thus, conventionally, there is no double-flowering variety among ornamental strawberry varieties. In addition, when the existing variety is used for ground cover applications, it is necessary to remove the decayed fruits over time.
本発明は、上述した実情に鑑み、純度の高いＦ１採種の効率化のために、雄性不稔を有するイチゴ交配母本を提供することを目的とする。
上記課題を解決するため鋭意研究を行った結果、フラガリア・イイヌマエ由来の細胞質とイチゴ栽培種の核ゲノムとの組合せを有するイチゴは、核遺伝子細胞質雄性不稔となることを見出し、本発明を完成するに至った。
すなわち、本発明は、以下を包含する。
（１）フラガリア・イイヌマエ（Ｆｒａｇａｒｉａ ｉｉｎｕｍａｅ）由来の細胞質を有し、且つ核遺伝子細胞質雄性不稔を有するイチゴ又はその後代。
（２）イチゴ栽培種の核ゲノムを有する、（１）記載のイチゴ又はその後代。
（３）イチゴ栽培種がフラガリアｘアナナッサ（Ｆｒａｇａｒｉａ ｘ ａｎａｎａｓｓａ）、又はフラガリアｘアナナッサとフラガリア・ニルゲレンシス（Ｆｒａｇａｒｉａ ｎｉｌｇｅｒｒｅｎｓｉｓ）との種間雑種である、（２）記載のイチゴ又はその後代。
（４）（１）~（３）のいずれか１記載のイチゴ又はその後代の植物体の一部。
（５）植物体の一部が種子である、（４）記載の植物体の一部。
（６）フラガリア・イイヌマエを種子親としてイチゴ栽培種と交配し、フラガリア・イイヌマエ由来の細胞質を有する種間雑種を作出する工程と、前記種間雑種を種子親としてイチゴ栽培種と連続戻し交配する工程とを含む、フラガリア・イイヌマエ由来の細胞質を有し、且つ核遺伝子細胞質雄性不稔を有するイチゴの作出方法。
（７）連続戻し交配工程前に、種間雑種を染色体倍加処理に供する工程をさらに含む、（６）記載の方法。
（８）イチゴ栽培種がフラガリアｘアナナッサ、又はフラガリアｘアナナッサとフラガリア・ニルゲレンシスとの種間雑種である、（７）記載の方法。
（９）（１）~（３）のいずれか１記載のイチゴ又はその後代を種子親とし、該イチゴ又はその後代と交配可能なイチゴ栽培種を花粉親として交配し、交配後の種子親から雑種第一代種子（Ｆ１種子）を採種する工程を含む、Ｆ１種子の作出方法。
（１０）（１）~（３）のいずれか１記載のイチゴ又はその後代と、有用形質を有するイチゴ品種を連続戻し交配する工程を含む、前記有用形質を有し、且つ核遺伝子細胞質雄性不稔を有するイチゴの作出方法。
本明細書は本願の優先権の基礎となる日本国特許出願番号２０１７−０４０８５１の開示内容を包含する。 In view of the above-described circumstances, an object of the present invention is to provide a strawberry mating mother having male sterility in order to improve the efficiency of F1 seeding with high purity.
As a result of diligent research to solve the above problems, it was found that a strawberry having a combination of the cytoplasm derived from Fragalia iinumae and the nuclear genome of a strawberry cultivar becomes a nuclear gene cytoplasmic male sterile and completed the present invention. It came to do.
That is, the present invention includes the following.
(1) A strawberry or progeny having cytoplasm derived from Fragaria iinumae and having a nuclear gene cytoplasmic male sterility.
(2) The strawberry or its progeny according to (1), which has a nuclear genome of a strawberry cultivated species.
(3) The strawberry or progeny thereof according to (2), wherein the strawberry cultivated species is Fragaria x ananassa, or an interspecific hybrid of Fragalia x ananassa and Fragalia nigerrensis (Fragaria nilgerrensis).
(4) A part of the strawberry according to any one of (1) to (3) or a plant of a progeny thereof.
(5) A part of the plant according to (4), wherein a part of the plant is a seed.
(6) Crossing with strawberry cultivated species using fragalia iinumae as a seed parent to produce an interspecific hybrid having cytoplasm derived from fragalia iinumae, and continuously backcrossing with strawberry cultivated species using the interspecific hybrid as a seed parent. And a method for producing a strawberry having a cytoplasm derived from Fragalia iinumae and having a nuclear gene cytoplasmic male sterility.
(7) The method according to (6), further comprising a step of subjecting the interspecific hybrid to a chromosome doubling treatment before the continuous backcrossing step.
(8) The method according to (7), wherein the strawberry cultivated species is Fragalia x Ananassa, or an interspecific hybrid of Fragalia x Ananassa and Fragalia nigerensis.
(9) A strawberry according to any one of (1) to (3) or a progeny thereof is used as a seed parent, and a strawberry cultivated species that can be crossed with the strawberry or its progeny is crossed as a pollen parent. A method for producing F1 seeds, comprising the step of seeding hybrid first-generation seeds (F1 seeds).
(10) The strawberry of any one of (1) to (3) or a progeny thereof and the step of continuously backcrossing a strawberry variety having a useful trait and having the above-mentioned useful trait and having a nuclear gene cytoplasmic male How to make strawberries with strawberries.
This specification includes the disclosure of Japanese Patent Application No. 2017-040851, which is the basis of the priority of the present application.
上記課題を解決するため鋭意研究を行った結果、フラガリア・イイヌマエ由来の細胞質とイチゴ栽培種の核ゲノムとの組合せを有するイチゴは、核遺伝子細胞質雄性不稔となることを見出し、本発明を完成するに至った。
すなわち、本発明は、以下を包含する。
（１）フラガリア・イイヌマエ（Ｆｒａｇａｒｉａ ｉｉｎｕｍａｅ）由来の細胞質を有し、且つ核遺伝子細胞質雄性不稔を有するイチゴ又はその後代。
（２）イチゴ栽培種の核ゲノムを有する、（１）記載のイチゴ又はその後代。
（３）イチゴ栽培種がフラガリアｘアナナッサ（Ｆｒａｇａｒｉａ ｘ ａｎａｎａｓｓａ）、又はフラガリアｘアナナッサとフラガリア・ニルゲレンシス（Ｆｒａｇａｒｉａ ｎｉｌｇｅｒｒｅｎｓｉｓ）との種間雑種である、（２）記載のイチゴ又はその後代。
（４）（１）~（３）のいずれか１記載のイチゴ又はその後代の植物体の一部。
（５）植物体の一部が種子である、（４）記載の植物体の一部。
（６）フラガリア・イイヌマエを種子親としてイチゴ栽培種と交配し、フラガリア・イイヌマエ由来の細胞質を有する種間雑種を作出する工程と、前記種間雑種を種子親としてイチゴ栽培種と連続戻し交配する工程とを含む、フラガリア・イイヌマエ由来の細胞質を有し、且つ核遺伝子細胞質雄性不稔を有するイチゴの作出方法。
（７）連続戻し交配工程前に、種間雑種を染色体倍加処理に供する工程をさらに含む、（６）記載の方法。
（８）イチゴ栽培種がフラガリアｘアナナッサ、又はフラガリアｘアナナッサとフラガリア・ニルゲレンシスとの種間雑種である、（７）記載の方法。
（９）（１）~（３）のいずれか１記載のイチゴ又はその後代を種子親とし、該イチゴ又はその後代と交配可能なイチゴ栽培種を花粉親として交配し、交配後の種子親から雑種第一代種子（Ｆ１種子）を採種する工程を含む、Ｆ１種子の作出方法。
（１０）（１）~（３）のいずれか１記載のイチゴ又はその後代と、有用形質を有するイチゴ品種を連続戻し交配する工程を含む、前記有用形質を有し、且つ核遺伝子細胞質雄性不稔を有するイチゴの作出方法。
本明細書は本願の優先権の基礎となる日本国特許出願番号２０１７−０４０８５１の開示内容を包含する。 In view of the above-described circumstances, an object of the present invention is to provide a strawberry mating mother having male sterility in order to improve the efficiency of F1 seeding with high purity.
As a result of diligent research to solve the above problems, it was found that a strawberry having a combination of the cytoplasm derived from Fragalia iinumae and the nuclear genome of a strawberry cultivar becomes a nuclear gene cytoplasmic male sterile and completed the present invention. It came to do.
That is, the present invention includes the following.
(1) A strawberry or progeny having cytoplasm derived from Fragaria iinumae and having a nuclear gene cytoplasmic male sterility.
(2) The strawberry or its progeny according to (1), which has a nuclear genome of a strawberry cultivated species.
(3) The strawberry or progeny thereof according to (2), wherein the strawberry cultivated species is Fragaria x ananassa, or an interspecific hybrid of Fragalia x ananassa and Fragalia nigerrensis (Fragaria nilgerrensis).
(4) A part of the strawberry according to any one of (1) to (3) or a plant of a progeny thereof.
(5) A part of the plant according to (4), wherein a part of the plant is a seed.
(6) Crossing with strawberry cultivated species using fragalia iinumae as a seed parent to produce an interspecific hybrid having cytoplasm derived from fragalia iinumae, and continuously backcrossing with strawberry cultivated species using the interspecific hybrid as a seed parent. And a method for producing a strawberry having a cytoplasm derived from Fragalia iinumae and having a nuclear gene cytoplasmic male sterility.
(7) The method according to (6), further comprising a step of subjecting the interspecific hybrid to a chromosome doubling treatment before the continuous backcrossing step.
(8) The method according to (7), wherein the strawberry cultivated species is Fragalia x Ananassa, or an interspecific hybrid of Fragalia x Ananassa and Fragalia nigerensis.
(9) A strawberry according to any one of (1) to (3) or a progeny thereof is used as a seed parent, and a strawberry cultivated species that can be crossed with the strawberry or its progeny is crossed as a pollen parent. A method for producing F1 seeds, comprising the step of seeding hybrid first-generation seeds (F1 seeds).
(10) The strawberry of any one of (1) to (3) or a progeny thereof and the step of continuously backcrossing a strawberry variety having a useful trait and having the above-mentioned useful trait and having a nuclear gene cytoplasmic male How to make strawberries with strawberries.
This specification includes the disclosure of Japanese Patent Application No. 2017-040851, which is the basis of the priority of the present application.
本発明に係る核遺伝子細胞質雄性不稔（又は細胞質雄性不稔）を有するイチゴ又はその後代は、２倍体野生種フラガリア・イイヌマエ由来の細胞質を有するものである。核遺伝子細胞質雄性不稔は、ミトコンドリアゲノム上に存在する特有のｏｒｆによって引き起こされ、母性遺伝する形質である。当該形質は、核ゲノムに存在する稔性回復遺伝子によって打ち消され、正常に花粉が形成される。
本発明は、フラガリア・イイヌマエの核ゲノムには稔性回復遺伝子が存在するが、フラガリア・イイヌマエ由来の細胞質とイチゴ栽培種の核ゲノムとの組合せを有するイチゴが核遺伝子細胞質雄性不稔となるという知見に基づくものである。
本発明に係る核遺伝子細胞質雄性不稔を有するイチゴが有する核ゲノムが由来するイチゴ栽培種としては、例えばフラガリアｘアナナッサ（８倍体栽培種）、フラガリアｘアナナッサと２倍体野生種フラガリア・ニルゲレンシスとの種間雑種（１０倍体栽培種；例えば品種「桃薫」）等が挙げられる。
また、本発明に係る核遺伝子細胞質雄性不稔を有するイチゴの後代とは、本発明に係る核遺伝子細胞質雄性不稔性を有するイチゴを種子親とし、該イチゴに交配可能なイチゴ品種を花粉親として交配することによって得られる、母性遺伝により雄性不稔性を引き継ぐ次世代及び次世代以降のイチゴを意味する。このような本発明に係る核遺伝子細胞質雄性不稔を有するイチゴに交配可能なイチゴ品種としては、例えば‘さちのか’、‘カレンベリー’、‘かおり野’、‘おおきみ’、‘桃薫’、‘久留米ＩＨ４号’、‘彩紅姫’等が挙げられる。
さらに、本発明に係る核遺伝子細胞質雄性不稔を有するイチゴ又はその後代の植物体の一部としては、例えば種子や花、葉、茎、根等の器官又は組織、或いは、これらの器官又は組織からの細胞若しくは細胞質が挙げられる。
本発明に係る核遺伝子細胞質雄性不稔を有するイチゴの作出方法を図１に示す。図１において、「ｉ」はフラガリア・イイヌマエのミトコンドリアゲノムを示し、「Ｉ」はフラガリア・イイヌマエの核の染色体を示し、「ａ」はフラガリアｘアナナッサ（Ｆ．ｘ ａｎａｎａｓｓａ）のミトコンドリアゲノムを示し、且つ「Ａ」はフラガリアｘアナナッサの核の染色体を示す。
当該作出方法では、先ず、フラガリア・イイヌマエ（２倍体）を種子親としてイチゴ栽培種と交配し、フラガリア・イイヌマエ由来の細胞質を有する種間雑種を作出する。
図１に示すように、例えば８倍体栽培種であるフラガリアｘアナナッサをイチゴ栽培種として交配すると、得られる種間雑種は５倍体であり、雌性稔性も花粉形成能も有しない。そこで、このような奇数倍数体の種間雑種が作出される場合には、連続戻し交配工程前に、当該種間雑種を例えばコルヒチンを用いた染色体倍加処理に供することで、雌性稔性及び花粉形成能のある偶数倍数体（図１では１０倍体）の種間雑種を作出する。
次いで、種間雑種を種子親として、種間雑種作出に使用したイチゴ栽培種又は他のイチゴ栽培種と連続戻し交配する。当該戻し交配を進めると、成熟花粉を形成しない核遺伝子細胞質雄性不稔を有するイチゴが出現する。換言すれば、当該成熟花粉を形成しない個体を、本発明に係る核遺伝子細胞質雄性不稔を有するイチゴとして選抜することができる。
また、作出した本発明に係る核遺伝子細胞質雄性不稔を有するイチゴ又はその後代を種子親とし、該イチゴ又はその後代と交配可能なイチゴ栽培種を花粉親として交配し、交配後の種子親からＦ１種子を採種することができる。
さらに、作出した本発明に係る核遺伝子細胞質雄性不稔を有するイチゴ又はその後代に、有用形質（例えば、果実品質若しくは四季成り性）を有するイチゴ品種を連続戻し交配することにより、後代として、当該有用形質を有し、且つ核遺伝子細胞質雄性不稔性を有する優良系統を得ることができる。このような有用形質を有するイチゴ品種としては、例えば‘三重母本１号’、‘Ａ８Ｓ４−１４７’等が挙げられる。当該核遺伝子細胞質雄性不稔性を有する優良系統は、Ｆ１種子を得るための種子親として使用することができる。
以上に説明する本発明に係る核遺伝子細胞質雄性不稔を有するイチゴは、以下の特徴を有する。
（１）従来技術によるイチゴのＦ１採種は人手による除雄と交配が行われており、高い人件費と自殖種子混入の危険性が伴う。一方、本発明に係る花粉形成の認められない核遺伝子細胞質雄性不稔を有するイチゴ系統を種子親として用いることにより、花粉親系統と混植し花粉媒介昆虫を放飼するのみで、低コスト且つ確実にＦ１種子を得ることができる。
（２）本発明に係る核遺伝子細胞質雄性不稔を有するイチゴは、雄ずいが花弁化したものであり、成熟花粉が形成されないペタロイド型の細胞質雄性不稔を有する。当該雄性不稔イチゴに栽培種を交配すると、多くの実生が成熟花粉を形成できない雄ずい形態異常を呈することから、連続戻し交配により既存の栽培種（有望自殖系統）を雄性不稔化できる。
（３）本発明に係る核遺伝子細胞質雄性不稔を有するイチゴは、フラガリア・イイヌマエの核ゲノム（稔性回復遺伝子）を持つ１０倍体種間雑種を交配することにより、雄ずいの形態が正常化し、花粉形成能が復活（稔性回復）した実生を作出することが可能である。従って、本発明に係る核遺伝子細胞質雄性不稔を有するイチゴを種子親とし、且つフラガリア・イイヌマエの稔性回復遺伝子を持つ系統を花粉親とした、効率的なＦ１採種技術が開発できる。
（４）雄ずいが花弁化する系統は八重咲きとなり、観賞用品種育成に利用できる。観賞性の高い八重化したイチゴ系統は、成熟花粉を形成しない雄性不稔であるため、他のイチゴ品種と混植をしない限り果実肥大せず、腐敗果がほとんど生じないため、適応場面は広い。 Strawberries or progenies having the nuclear gene cytoplasmic male sterility (or cytoplasmic male sterility) according to the present invention have a cytoplasm derived from a diploid wild species, Fragalia iinumae. The nuclear gene cytoplasmic male sterility is a maternally inherited trait caused by a unique orf present on the mitochondrial genome. The trait is counteracted by the fertility restoration gene present in the nuclear genome, and pollen is normally formed.
According to the present invention, there is a fertility-recovering gene in the nuclear genome of Fragalia iinumae, but a strawberry having a combination of cytoplasm derived from Fragalia iinumae and the nuclear genome of a strawberry cultivar is said to be a nuclear gene cytoplasmic male sterile. Based on knowledge.
Strawberry cultivars from which the nuclear genome possessed by the strawberry having nuclear gene cytoplasmic male sterility according to the present invention are derived, for example, Fragalia x Ananassa (octaploid cultivar), Fragalia x Ananassa and diploid wild species Fragalia nilgrensis And interspecific hybrids (deploid cultivated species; for example, cultivar “Momochi”) and the like.
The progeny of a strawberry having a nuclear gene cytoplasmic male sterility according to the present invention refers to a strawberry having a nuclear gene cytoplasmic male sterility according to the present invention as a seed parent and a strawberry variety that can be crossed with the strawberry as a pollen parent. It means the next generation and subsequent generations of strawberries that inherit male sterility by maternal inheritance. Strawberry varieties that can be crossed with strawberry having nuclear gene cytoplasmic male sterility according to the present invention include, for example, 'Sachinoka', 'Cullenberry', 'Kaorino', 'Ookimi', 'Momochi' , “Kurume IH4”, “Saikuhime” and the like.
Furthermore, as a part of a strawberry having a nuclear gene cytoplasmic male sterility or a progeny plant thereof according to the present invention, for example, organs or tissues such as seeds, flowers, leaves, stems, roots, or these organs or tissues Cell or cytoplasm.
A method for producing a strawberry having a nuclear gene cytoplasmic male sterility according to the present invention is shown in FIG. In FIG. 1, “i” represents the mitochondrial genome of Fraglia iinumae, “I” represents the chromosome of the nucleus of Fraglia iinumae, “a” represents the mitochondrial genome of F. x ananassa, "A" indicates the chromosome of the nucleus of Fragalia x Ananassa.
In the production method, first, a strawberry cultivated species is crossed using fragalia iinumae (diploid) as a seed parent to produce an interspecific hybrid having cytoplasm derived from fragalia iinumae.
As shown in FIG. 1, for example, when Fragalia x Ananassa, which is an octaploid cultivar, is crossed as a strawberry cultivar, the interspecific hybrid obtained is a pentaploid and has neither female fertility nor pollen-forming ability. Therefore, when such an odd polyploid interspecific hybrid is created, before the continuous backcrossing step, the interspecific hybrid is subjected to chromosome doubling treatment using, for example, colchicine, so that female fertility and pollen An interspecific hybrid of even-numbered polyploids (deploids in FIG. 1) having a forming ability is produced.
Next, using the interspecific hybrid as a seed parent, continuous backcrossing is performed with the strawberry cultivated species or other strawberry cultivated species used for interspecific hybrid production. As the backcrossing proceeds, strawberries with nuclear gene cytoplasmic male sterility that do not form mature pollen appear. In other words, individuals that do not form mature pollen can be selected as strawberries having the nuclear gene cytoplasmic male sterility according to the present invention.
In addition, the strawberry having a nuclear gene cytoplasmic male sterility according to the present invention or its progeny is used as a seed parent, and the strawberry cultivated species that can be crossed with the strawberry or its progeny is crossed as a pollen parent, from the seed parent after the crossing F1 seeds can be collected.
Furthermore, the progeny can be obtained by continuously backcrossing a strawberry variety having a useful trait (for example, fruit quality or seasonality) to a strawberry having a nuclear gene cytoplasmic male sterility according to the present invention or its progeny. An excellent line having useful traits and having a nuclear gene cytoplasmic male sterility can be obtained. Examples of strawberry varieties having such useful traits include “Mie Motomoto No. 1” and “A8S4-147”. The excellent line having the nuclear gene cytoplasmic male sterility can be used as a seed parent for obtaining F1 seeds.
The strawberry having the nuclear gene cytoplasmic male sterility according to the present invention described above has the following characteristics.
(1) F1 seeding of strawberries according to the prior art is manually sterilized and mated, which is associated with high labor costs and the risk of contamination of self-grown seeds. On the other hand, by using a strawberry line having a nuclear gene cytoplasmic male sterility in which no pollen formation according to the present invention is used as a seed parent, it is possible to reduce the cost and reliability by simply mixing the pollen parent line and releasing pollen-borne insects. F1 seeds can be obtained.
(2) A strawberry having a nuclear gene cytoplasmic male sterility according to the present invention is a petalized stamen and has a petaloid cytoplasmic male sterility in which mature pollen is not formed. When cultivated species are mated to the male sterile strawberries, many seedlings exhibit stamen morphology abnormalities that cannot form mature pollen, so existing cultivated species (promising inbred lines) can be sterilized by continuous backcrossing .
(3) Strawberry having a nuclear gene cytoplasmic male sterility according to the present invention has normal stamen morphology by crossing a haploid interspecific hybrid having a nuclear genome (fertility recovery gene) of fragalia iinumae It is possible to produce seedlings that have revived (recovered fertility) pollen formation ability. Therefore, it is possible to develop an efficient F1 seeding technique in which a strawberry having a nuclear gene cytoplasmic male sterility according to the present invention is used as a seed parent and a line having a fertility recovery gene of fragalia iinumae is used as a pollen parent.
(4) The line where the stamens petal will be in double bloom and can be used for breeding ornamental varieties. The highly ornamental octuplicated strawberry line is a male sterility that does not form mature pollen. Therefore, unless it is mixed with other strawberry varieties, the fruit does not enlarge and there is almost no spoilage.
本発明は、フラガリア・イイヌマエの核ゲノムには稔性回復遺伝子が存在するが、フラガリア・イイヌマエ由来の細胞質とイチゴ栽培種の核ゲノムとの組合せを有するイチゴが核遺伝子細胞質雄性不稔となるという知見に基づくものである。
本発明に係る核遺伝子細胞質雄性不稔を有するイチゴが有する核ゲノムが由来するイチゴ栽培種としては、例えばフラガリアｘアナナッサ（８倍体栽培種）、フラガリアｘアナナッサと２倍体野生種フラガリア・ニルゲレンシスとの種間雑種（１０倍体栽培種；例えば品種「桃薫」）等が挙げられる。
また、本発明に係る核遺伝子細胞質雄性不稔を有するイチゴの後代とは、本発明に係る核遺伝子細胞質雄性不稔性を有するイチゴを種子親とし、該イチゴに交配可能なイチゴ品種を花粉親として交配することによって得られる、母性遺伝により雄性不稔性を引き継ぐ次世代及び次世代以降のイチゴを意味する。このような本発明に係る核遺伝子細胞質雄性不稔を有するイチゴに交配可能なイチゴ品種としては、例えば‘さちのか’、‘カレンベリー’、‘かおり野’、‘おおきみ’、‘桃薫’、‘久留米ＩＨ４号’、‘彩紅姫’等が挙げられる。
さらに、本発明に係る核遺伝子細胞質雄性不稔を有するイチゴ又はその後代の植物体の一部としては、例えば種子や花、葉、茎、根等の器官又は組織、或いは、これらの器官又は組織からの細胞若しくは細胞質が挙げられる。
本発明に係る核遺伝子細胞質雄性不稔を有するイチゴの作出方法を図１に示す。図１において、「ｉ」はフラガリア・イイヌマエのミトコンドリアゲノムを示し、「Ｉ」はフラガリア・イイヌマエの核の染色体を示し、「ａ」はフラガリアｘアナナッサ（Ｆ．ｘ ａｎａｎａｓｓａ）のミトコンドリアゲノムを示し、且つ「Ａ」はフラガリアｘアナナッサの核の染色体を示す。
当該作出方法では、先ず、フラガリア・イイヌマエ（２倍体）を種子親としてイチゴ栽培種と交配し、フラガリア・イイヌマエ由来の細胞質を有する種間雑種を作出する。
図１に示すように、例えば８倍体栽培種であるフラガリアｘアナナッサをイチゴ栽培種として交配すると、得られる種間雑種は５倍体であり、雌性稔性も花粉形成能も有しない。そこで、このような奇数倍数体の種間雑種が作出される場合には、連続戻し交配工程前に、当該種間雑種を例えばコルヒチンを用いた染色体倍加処理に供することで、雌性稔性及び花粉形成能のある偶数倍数体（図１では１０倍体）の種間雑種を作出する。
次いで、種間雑種を種子親として、種間雑種作出に使用したイチゴ栽培種又は他のイチゴ栽培種と連続戻し交配する。当該戻し交配を進めると、成熟花粉を形成しない核遺伝子細胞質雄性不稔を有するイチゴが出現する。換言すれば、当該成熟花粉を形成しない個体を、本発明に係る核遺伝子細胞質雄性不稔を有するイチゴとして選抜することができる。
また、作出した本発明に係る核遺伝子細胞質雄性不稔を有するイチゴ又はその後代を種子親とし、該イチゴ又はその後代と交配可能なイチゴ栽培種を花粉親として交配し、交配後の種子親からＦ１種子を採種することができる。
さらに、作出した本発明に係る核遺伝子細胞質雄性不稔を有するイチゴ又はその後代に、有用形質（例えば、果実品質若しくは四季成り性）を有するイチゴ品種を連続戻し交配することにより、後代として、当該有用形質を有し、且つ核遺伝子細胞質雄性不稔性を有する優良系統を得ることができる。このような有用形質を有するイチゴ品種としては、例えば‘三重母本１号’、‘Ａ８Ｓ４−１４７’等が挙げられる。当該核遺伝子細胞質雄性不稔性を有する優良系統は、Ｆ１種子を得るための種子親として使用することができる。
以上に説明する本発明に係る核遺伝子細胞質雄性不稔を有するイチゴは、以下の特徴を有する。
（１）従来技術によるイチゴのＦ１採種は人手による除雄と交配が行われており、高い人件費と自殖種子混入の危険性が伴う。一方、本発明に係る花粉形成の認められない核遺伝子細胞質雄性不稔を有するイチゴ系統を種子親として用いることにより、花粉親系統と混植し花粉媒介昆虫を放飼するのみで、低コスト且つ確実にＦ１種子を得ることができる。
（２）本発明に係る核遺伝子細胞質雄性不稔を有するイチゴは、雄ずいが花弁化したものであり、成熟花粉が形成されないペタロイド型の細胞質雄性不稔を有する。当該雄性不稔イチゴに栽培種を交配すると、多くの実生が成熟花粉を形成できない雄ずい形態異常を呈することから、連続戻し交配により既存の栽培種（有望自殖系統）を雄性不稔化できる。
（３）本発明に係る核遺伝子細胞質雄性不稔を有するイチゴは、フラガリア・イイヌマエの核ゲノム（稔性回復遺伝子）を持つ１０倍体種間雑種を交配することにより、雄ずいの形態が正常化し、花粉形成能が復活（稔性回復）した実生を作出することが可能である。従って、本発明に係る核遺伝子細胞質雄性不稔を有するイチゴを種子親とし、且つフラガリア・イイヌマエの稔性回復遺伝子を持つ系統を花粉親とした、効率的なＦ１採種技術が開発できる。
（４）雄ずいが花弁化する系統は八重咲きとなり、観賞用品種育成に利用できる。観賞性の高い八重化したイチゴ系統は、成熟花粉を形成しない雄性不稔であるため、他のイチゴ品種と混植をしない限り果実肥大せず、腐敗果がほとんど生じないため、適応場面は広い。 Strawberries or progenies having the nuclear gene cytoplasmic male sterility (or cytoplasmic male sterility) according to the present invention have a cytoplasm derived from a diploid wild species, Fragalia iinumae. The nuclear gene cytoplasmic male sterility is a maternally inherited trait caused by a unique orf present on the mitochondrial genome. The trait is counteracted by the fertility restoration gene present in the nuclear genome, and pollen is normally formed.
According to the present invention, there is a fertility-recovering gene in the nuclear genome of Fragalia iinumae, but a strawberry having a combination of cytoplasm derived from Fragalia iinumae and the nuclear genome of a strawberry cultivar is said to be a nuclear gene cytoplasmic male sterile. Based on knowledge.
Strawberry cultivars from which the nuclear genome possessed by the strawberry having nuclear gene cytoplasmic male sterility according to the present invention are derived, for example, Fragalia x Ananassa (octaploid cultivar), Fragalia x Ananassa and diploid wild species Fragalia nilgrensis And interspecific hybrids (deploid cultivated species; for example, cultivar “Momochi”) and the like.
The progeny of a strawberry having a nuclear gene cytoplasmic male sterility according to the present invention refers to a strawberry having a nuclear gene cytoplasmic male sterility according to the present invention as a seed parent and a strawberry variety that can be crossed with the strawberry as a pollen parent. It means the next generation and subsequent generations of strawberries that inherit male sterility by maternal inheritance. Strawberry varieties that can be crossed with strawberry having nuclear gene cytoplasmic male sterility according to the present invention include, for example, 'Sachinoka', 'Cullenberry', 'Kaorino', 'Ookimi', 'Momochi' , “Kurume IH4”, “Saikuhime” and the like.
Furthermore, as a part of a strawberry having a nuclear gene cytoplasmic male sterility or a progeny plant thereof according to the present invention, for example, organs or tissues such as seeds, flowers, leaves, stems, roots, or these organs or tissues Cell or cytoplasm.
A method for producing a strawberry having a nuclear gene cytoplasmic male sterility according to the present invention is shown in FIG. In FIG. 1, “i” represents the mitochondrial genome of Fraglia iinumae, “I” represents the chromosome of the nucleus of Fraglia iinumae, “a” represents the mitochondrial genome of F. x ananassa, "A" indicates the chromosome of the nucleus of Fragalia x Ananassa.
In the production method, first, a strawberry cultivated species is crossed using fragalia iinumae (diploid) as a seed parent to produce an interspecific hybrid having cytoplasm derived from fragalia iinumae.
As shown in FIG. 1, for example, when Fragalia x Ananassa, which is an octaploid cultivar, is crossed as a strawberry cultivar, the interspecific hybrid obtained is a pentaploid and has neither female fertility nor pollen-forming ability. Therefore, when such an odd polyploid interspecific hybrid is created, before the continuous backcrossing step, the interspecific hybrid is subjected to chromosome doubling treatment using, for example, colchicine, so that female fertility and pollen An interspecific hybrid of even-numbered polyploids (deploids in FIG. 1) having a forming ability is produced.
Next, using the interspecific hybrid as a seed parent, continuous backcrossing is performed with the strawberry cultivated species or other strawberry cultivated species used for interspecific hybrid production. As the backcrossing proceeds, strawberries with nuclear gene cytoplasmic male sterility that do not form mature pollen appear. In other words, individuals that do not form mature pollen can be selected as strawberries having the nuclear gene cytoplasmic male sterility according to the present invention.
In addition, the strawberry having a nuclear gene cytoplasmic male sterility according to the present invention or its progeny is used as a seed parent, and the strawberry cultivated species that can be crossed with the strawberry or its progeny is crossed as a pollen parent, from the seed parent after the crossing F1 seeds can be collected.
Furthermore, the progeny can be obtained by continuously backcrossing a strawberry variety having a useful trait (for example, fruit quality or seasonality) to a strawberry having a nuclear gene cytoplasmic male sterility according to the present invention or its progeny. An excellent line having useful traits and having a nuclear gene cytoplasmic male sterility can be obtained. Examples of strawberry varieties having such useful traits include “Mie Motomoto No. 1” and “A8S4-147”. The excellent line having the nuclear gene cytoplasmic male sterility can be used as a seed parent for obtaining F1 seeds.
The strawberry having the nuclear gene cytoplasmic male sterility according to the present invention described above has the following characteristics.
(1) F1 seeding of strawberries according to the prior art is manually sterilized and mated, which is associated with high labor costs and the risk of contamination of self-grown seeds. On the other hand, by using a strawberry line having a nuclear gene cytoplasmic male sterility in which no pollen formation according to the present invention is used as a seed parent, it is possible to reduce the cost and reliability by simply mixing the pollen parent line and releasing pollen-borne insects. F1 seeds can be obtained.
(2) A strawberry having a nuclear gene cytoplasmic male sterility according to the present invention is a petalized stamen and has a petaloid cytoplasmic male sterility in which mature pollen is not formed. When cultivated species are mated to the male sterile strawberries, many seedlings exhibit stamen morphology abnormalities that cannot form mature pollen, so existing cultivated species (promising inbred lines) can be sterilized by continuous backcrossing .
(3) Strawberry having a nuclear gene cytoplasmic male sterility according to the present invention has normal stamen morphology by crossing a haploid interspecific hybrid having a nuclear genome (fertility recovery gene) of fragalia iinumae It is possible to produce seedlings that have revived (recovered fertility) pollen formation ability. Therefore, it is possible to develop an efficient F1 seeding technique in which a strawberry having a nuclear gene cytoplasmic male sterility according to the present invention is used as a seed parent and a line having a fertility recovery gene of fragalia iinumae is used as a pollen parent.
(4) The line where the stamens petal will be in double bloom and can be used for breeding ornamental varieties. The highly ornamental octuplicated strawberry line is a male sterility that does not form mature pollen. Therefore, unless it is mixed with other strawberry varieties, the fruit does not enlarge and there is almost no spoilage.
以下、実施例を用いて本発明をより詳細に説明するが、本発明の技術的範囲はこれら実施例に限定されるものではない。
１．材料及び方法
２００８年に２倍体野生種フラガリア・イイヌマエ（Ｆ．ｉｉｎｕｍａｅ）に８倍体栽培品種‘カレンベリー’（イチゴ栽培種フラガリアｘアナナッサ）を交配することにより、不稔５倍体種間雑種０８ＩＫ５８０４を得た。２００９年に０８ＩＫ５８０４をコルヒチンで倍加処理することによって作出した、雌性稔性及び花粉形成能を有する１０倍体種間雑種０８ＩＫ５８０４−０１を種子親として供試した。花粉親としては８倍体イチゴ栽培種のほか、１０倍体種間雑種０８ＳｃｈＩ２８−０１（さちのか×Ｆ．ｉｉｎｕｍａｅ）を供試した。野生種（Ｆ．ｉｉｎｕｍａｅ）の細胞質を有する０８ＩＫ５８０４−０１に、イチゴ栽培種を連続戻し交配することによって、核遺伝子細胞質雄性不稔系統の作出を試みた。
２．結果及び考察
０８ＩＫ５８０４−０１に、若しくは０８ＩＫ５８０４−０１に０８ＳｃｈＩ２８−０１を交配したＤＨ１０２０−１７に、イチゴ栽培種を連続的に戻し交配することによって、雄ずいが花弁化して正常な成熟花粉を形成できない雄ずい形態異常（ペタロイド型雄性不稔）個体が出現した（図２）。雄ずい形態異常個体は、特定の系譜のみではなく、複数の系譜で出現した。
得られた雄ずい形態異常系統の内、下記の表１に示す２１系統にイチゴ栽培種を交配したところ、実生には未熟と推測される雄ずいから完全に花弁化した雄ずいまで、異常化程度の異なる雄ずいを持つ個体が出現した（図３）が、多くの実生（８６％）が成熟花粉を形成できない雄ずい形態異常を呈する（表１）ことから、この形質の遺伝には細胞質が関与しており、連続戻し交配によって既存の栽培種（有望自殖系統）を雄ずい形態異常（雄性不稔）化できると考えられる。
この雄ずい形態異常系統に、Ｆ．ｉｉｎｕｍａｅの核ゲノムを持つ１０倍体種間雑種を交配したところ、雄ずいの形態異常個体の出現率は１．５％であり、ほとんどの実生で雄ずい形態の正常化が認められた（表２）。雄ずい形態異常系統にイチゴ栽培種を交配した場合には、多くが雄ずい形態異常となり、Ｆ．ｉｉｎｕｍａｅの核ゲノムを持つ種間雑種を交配した場合にはほとんどが正常化することから、Ｆ．ｉｉｎｕｍａｅの核ゲノムには回復遺伝子が存在し、花粉形成能が復活（稔性回復）した実生を作出することが可能であると考えられる。
雄ずい形態異常系統では雄ずいが花弁化するため、白色花弁で八重咲きを呈する系統が存在する。それらに赤色の花弁を有するイチゴ栽培種‘彩紅姫’や‘久留米ＩＨ４号’を交配することによって、赤色花弁を有する八重咲き系統の作出が可能であった。現在までに様々な花器形態、花弁色のイチゴ系統が容易に作出されている（図４）ことから、雄ずい形態異常系統は、観賞用イチゴの育種素材としても有用であると考えられる。
EXAMPLES Hereinafter, although this invention is demonstrated in detail using an Example, the technical scope of this invention is not limited to these Examples.
1. Materials and Methods By crossing the haploid cultivar 'Callenberry' (strawberry cultivar Fragalia x Ananassa) to the diploid wild species F. iinumae in 2008, between sterile pentaploid species Hybrid 08IK5804 was obtained. A 10-fold interspecific hybrid 08IK5804-01 produced by doubling treatment of 08IK5804 with colchicine in 2009 and having female fertility and pollen-forming ability was used as a seed parent. As a parent of pollen, in addition to the cultivar of octaploid strawberry, 10-ploid interspecific hybrid 08SchI28-01 (Sachinoka × F. Iinumae) was used. An attempt was made to create a nuclear gene cytoplasmic male sterile line by continuously backcrossing a strawberry cultivar to 08IK5804-01 having the cytoplasm of a wild species (F. iinumae).
2. Results and Discussion By continuously backcrossing strawberry cultivars to 08IK5804-01 or DH1020-17 crossed with 08IK5804-01 and 08SchI28-01, the stamens petalize and cannot form normal mature pollen A stamen morphological abnormality (petaloid male sterility) individuals appeared (FIG. 2). Individuals with abnormal stamens appeared not only in specific genealogy but also in multiple genealogy.
Strawberry cultivars were crossed to 21 strains shown in Table 1 below among the obtained stamen morphologically abnormal lines, and abnormalities were observed from stamens presumed to be immature to seedlings to completely petalized stamens. Individuals with stamens with different degrees appeared (Fig. 3), but many seedlings (86%) exhibit stamen abnormalities that cannot form mature pollen (Table 1). It is thought that existing cultivated species (promising inbred lines) can be made stamen abnormal (male sterility) by continuous backcrossing.
To this stamen morphological abnormal line, When hybrids of 10-ploid hybrids with the nuclear genome of iinumae were crossed, the appearance rate of stamen abnormalities was 1.5%, and normalization of stamens was observed in most seedlings (Table). 2). When strawberry cultivars are crossed to stamen morphologically abnormal lines, many stamen morphologies are abnormal. When interspecific hybrids having the nuclear genome of iinumae are crossed, most of them are normalized. There is a recovery gene in the nuclear genome of iinumae, and it is considered possible to produce seedlings whose pollen-forming ability has been restored (recovery of fertility).
Since stamens petalize in stamen morphologically abnormal lines, there are lines with white petals and double blooms. By mating them with strawberry cultivars 'Saikohime' and 'Kurume IH4' with red petals, it was possible to produce double-flowered lines with red petals. Various vase forms and petal-colored strawberry lines have been easily created so far (FIG. 4), and therefore, the stamen-shaped abnormal lines are considered useful as breeding materials for ornamental strawberries.
１．材料及び方法
２００８年に２倍体野生種フラガリア・イイヌマエ（Ｆ．ｉｉｎｕｍａｅ）に８倍体栽培品種‘カレンベリー’（イチゴ栽培種フラガリアｘアナナッサ）を交配することにより、不稔５倍体種間雑種０８ＩＫ５８０４を得た。２００９年に０８ＩＫ５８０４をコルヒチンで倍加処理することによって作出した、雌性稔性及び花粉形成能を有する１０倍体種間雑種０８ＩＫ５８０４−０１を種子親として供試した。花粉親としては８倍体イチゴ栽培種のほか、１０倍体種間雑種０８ＳｃｈＩ２８−０１（さちのか×Ｆ．ｉｉｎｕｍａｅ）を供試した。野生種（Ｆ．ｉｉｎｕｍａｅ）の細胞質を有する０８ＩＫ５８０４−０１に、イチゴ栽培種を連続戻し交配することによって、核遺伝子細胞質雄性不稔系統の作出を試みた。
２．結果及び考察
０８ＩＫ５８０４−０１に、若しくは０８ＩＫ５８０４−０１に０８ＳｃｈＩ２８−０１を交配したＤＨ１０２０−１７に、イチゴ栽培種を連続的に戻し交配することによって、雄ずいが花弁化して正常な成熟花粉を形成できない雄ずい形態異常（ペタロイド型雄性不稔）個体が出現した（図２）。雄ずい形態異常個体は、特定の系譜のみではなく、複数の系譜で出現した。
得られた雄ずい形態異常系統の内、下記の表１に示す２１系統にイチゴ栽培種を交配したところ、実生には未熟と推測される雄ずいから完全に花弁化した雄ずいまで、異常化程度の異なる雄ずいを持つ個体が出現した（図３）が、多くの実生（８６％）が成熟花粉を形成できない雄ずい形態異常を呈する（表１）ことから、この形質の遺伝には細胞質が関与しており、連続戻し交配によって既存の栽培種（有望自殖系統）を雄ずい形態異常（雄性不稔）化できると考えられる。
1. Materials and Methods By crossing the haploid cultivar 'Callenberry' (strawberry cultivar Fragalia x Ananassa) to the diploid wild species F. iinumae in 2008, between sterile pentaploid species Hybrid 08IK5804 was obtained. A 10-fold interspecific hybrid 08IK5804-01 produced by doubling treatment of 08IK5804 with colchicine in 2009 and having female fertility and pollen-forming ability was used as a seed parent. As a parent of pollen, in addition to the cultivar of octaploid strawberry, 10-ploid interspecific hybrid 08SchI28-01 (Sachinoka × F. Iinumae) was used. An attempt was made to create a nuclear gene cytoplasmic male sterile line by continuously backcrossing a strawberry cultivar to 08IK5804-01 having the cytoplasm of a wild species (F. iinumae).
2. Results and Discussion By continuously backcrossing strawberry cultivars to 08IK5804-01 or DH1020-17 crossed with 08IK5804-01 and 08SchI28-01, the stamens petalize and cannot form normal mature pollen A stamen morphological abnormality (petaloid male sterility) individuals appeared (FIG. 2). Individuals with abnormal stamens appeared not only in specific genealogy but also in multiple genealogy.
Strawberry cultivars were crossed to 21 strains shown in Table 1 below among the obtained stamen morphologically abnormal lines, and abnormalities were observed from stamens presumed to be immature to seedlings to completely petalized stamens. Individuals with stamens with different degrees appeared (Fig. 3), but many seedlings (86%) exhibit stamen abnormalities that cannot form mature pollen (Table 1). It is thought that existing cultivated species (promising inbred lines) can be made stamen abnormal (male sterility) by continuous backcrossing.
本発明に係る核遺伝子細胞質雄性不稔を有するイチゴを種子親に用いることによって、効率的で且つ純度の高いＦ１種子の採種技術確立に貢献する。また、本発明に係る核遺伝子細胞質雄性不稔を有するイチゴの雄ずいには程度の異なる形態異常（花弁化）が認められ、雄ずいが花弁化した雄性不稔（ペタロイド型雄性不稔）イチゴは、鑑賞価値の高い八重咲きイチゴの作出に利用可能である。
本明細書で引用した全ての刊行物、特許及び特許出願はそのまま引用により本明細書に組み入れられるものとする。 By using a strawberry having a nuclear gene cytoplasmic male sterility according to the present invention as a seed parent, it contributes to the establishment of an efficient and highly pure F1 seed collection technique. In addition, strawberry stamens with nuclear gene cytoplasmic male sterility according to the present invention have male sterility (petaloid male sterility) strawberry with morphological abnormalities (petalization) of different degrees and stamen petals. Can be used to produce double-flowered strawberries with high appreciation.
All publications, patents and patent applications cited herein are hereby incorporated by reference in their entirety.
本明細書で引用した全ての刊行物、特許及び特許出願はそのまま引用により本明細書に組み入れられるものとする。 By using a strawberry having a nuclear gene cytoplasmic male sterility according to the present invention as a seed parent, it contributes to the establishment of an efficient and highly pure F1 seed collection technique. In addition, strawberry stamens with nuclear gene cytoplasmic male sterility according to the present invention have male sterility (petaloid male sterility) strawberry with morphological abnormalities (petalization) of different degrees and stamen petals. Can be used to produce double-flowered strawberries with high appreciation.
All publications, patents and patent applications cited herein are hereby incorporated by reference in their entirety.
Claims (10)
- フラガリア・イイヌマエ（Ｆｒａｇａｒｉａ ｉｉｎｕｍａｅ）由来の細胞質を有し、且つ核遺伝子細胞質雄性不稔を有するイチゴ又はその後代。 Strawberry or its progeny having cytoplasm derived from Fragaria iinumae and having a male sterility of the nuclear gene cytoplasm.
- イチゴ栽培種の核ゲノムを有する、請求項１記載のイチゴ又はその後代。 The strawberry or its progeny according to claim 1, which has a nuclear genome of a strawberry cultivated species.
- イチゴ栽培種がフラガリアｘアナナッサ（Ｆｒａｇａｒｉａ ｘ ａｎａｎａｓｓａ）、又はフラガリアｘアナナッサとフラガリア・ニルゲレンシス（Ｆｒａｇａｒｉａ ｎｉｌｇｅｒｒｅｎｓｉｓ）との種間雑種である、請求項２記載のイチゴ又はその後代。 The strawberry or its progeny according to claim 2, wherein the strawberry cultivated species is Fragaria x ananassa, or an interspecific hybrid of Fragalia x ananassa and Fragalia nigerrensis.
- 請求項１~３のいずれか１項記載のイチゴ又はその後代の植物体の一部。 A part of the strawberry according to any one of claims 1 to 3 or a plant of its progeny.
- 植物体の一部が種子である、請求項４記載の植物体の一部。 The plant body part according to claim 4, wherein the plant body part is a seed.
- フラガリア・イイヌマエを種子親としてイチゴ栽培種と交配し、フラガリア・イイヌマエ由来の細胞質を有する種間雑種を作出する工程と、
前記種間雑種を種子親としてイチゴ栽培種と連続戻し交配する工程と、
を含む、フラガリア・イイヌマエ由来の細胞質を有し、且つ核遺伝子細胞質雄性不稔を有するイチゴの作出方法。 Crossing with strawberry cultivated species using fragalia iinumae as a seed parent, and producing an interspecific hybrid having cytoplasm derived from fragalia iinumae,
A step of continuously backcrossing with a strawberry cultivated seed as a seed parent of the interspecific hybrid;
A method for producing a strawberry having a cytoplasm derived from fragalia iinumae and having a male sterility of a nuclear gene cytoplasm. - 連続戻し交配工程前に、種間雑種を染色体倍加処理に供する工程をさらに含む、請求項６記載の方法。 The method according to claim 6, further comprising a step of subjecting the interspecific hybrid to a chromosome doubling treatment before the continuous backcrossing step.
- イチゴ栽培種がフラガリアｘアナナッサ、又はフラガリアｘアナナッサとフラガリア・ニルゲレンシスとの種間雑種である、請求項７記載の方法。 The method according to claim 7, wherein the strawberry cultivated species is Fragalia x Ananassa, or Interspecific hybrid of Fragalia x Ananassa and Fragalia nigerensis.
- 請求項１~３のいずれか１項記載のイチゴ又はその後代を種子親とし、該イチゴ又はその後代と交配可能なイチゴ栽培種を花粉親として交配し、交配後の種子親から雑種第一代種子を採種する工程を含む、雑種第一代種子の作出方法。 A strawberry or its progeny according to any one of claims 1 to 3 is used as a seed parent, a strawberry cultivated species that can be crossed with the strawberry or its progeny is crossed as a pollen parent, and a hybrid first from the seed parent after the crossing A method for producing hybrid primary seeds, comprising a step of seeding.
- 請求項１~３のいずれか１項記載のイチゴ又はその後代と、有用形質を有するイチゴ品種を連続戻し交配する工程を含む、前記有用形質を有し、且つ核遺伝子細胞質雄性不稔を有するイチゴの作出方法。 A strawberry having the useful trait and having a nuclear gene cytoplasmic male sterility, comprising the step of continuously backcrossing the strawberry of any one of claims 1 to 3 or its progeny and a strawberry variety having the useful trait How to make
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| JP2017040851A JP6833205B2 (en) | 2017-03-03 | 2017-03-03 | Strawberries with cytoplasm derived from Fragaria iinumae and with nuclear gene cytoplasmic male sterility (Gene-Cytoplasmic male sterility) |
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| CN111387049A (en) * | 2020-04-26 | 2020-07-10 | 贵州省园艺研究所(贵州省园艺工程技术研究中心) | Breeding method and application of early-maturing honey peach flavor type strawberries |
Citations (1)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| US5585540A (en) * | 1994-03-31 | 1996-12-17 | Sandoz Ltd. | Fragaria plants and seeds |
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| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| US5585540A (en) * | 1994-03-31 | 1996-12-17 | Sandoz Ltd. | Fragaria plants and seeds |
Non-Patent Citations (3)
| Title |
|---|
| ASHMAN TIA-LYNN ET AL.: "Multilocus Sex Determination Revealed in Two Populations of Gynodioecious Wild Strawberry, Fragaria vesca subsp. bracteata", G3 ; GENES, GENOMES, GENETICS, vol. 5, no. 12, 19 October 2015 (2015-10-19), pages 2759 - 2773, XP055538947, Retrieved from the Internet <URL:doi:10.1534/g3.115.023358> * |
| NOGUCHI YUJI ET AL.: "Interspecific Hybrids Originated from Crossing Asian Wild Strawberries (Fragaria nilgerrensis and F. iinumae) to F. xananassa", HORTSCIENCE, vol. 32, no. 3, June 1997 (1997-06-01), pages 438 - 439, XP055538949 * |
| vol. 11, no. 1, 2012, pages 112 * |
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| CN111387049A (en) * | 2020-04-26 | 2020-07-10 | 贵州省园艺研究所(贵州省园艺工程技术研究中心) | Breeding method and application of early-maturing honey peach flavor type strawberries |
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