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Frontiers of Agricultural Science and Engineering

ISSN 2095-7505

ISSN 2095-977X(Online)

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Front. Agr. Sci. Eng.    2021, Vol. 8 Issue (2) : 302-313    https://doi.org/10.15302/J-FASE-2021385
RESEARCH ARTICLE
UNREDUCED MEGAGAMETOPHYTE FORMATION VIA SECOND DIVISION RESTITUTION CONTRIBUTES TO TETRAPLOID PRODUCTION IN INTERPLOIDY CROSSES WITH ‘ORAH’ MANDARIN (CITRUS RETICULATA)
Qiangming XIA1, Wei WANG1, Kaidong XIE1(), Xiaomeng WU1, Xiuxin DENG1, Jude W. GROSSER2, Wenwu GUO1()
1. Key Laboratory of Horticultural Plant Biology (Ministry of Education), College of Horticulture and Forestry Sciences, Huazhong Agricultural University, Wuhan 430070, China.
2. Citrus Research and Education Center, University of Florida/IFAS, Lake Alfred, USA.
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Abstract

• In addition to triploid progeny, tetraploid hybrids derived from the fertilization of 2n megagametophytes are frequently regenerated from 2x × 4x crosses that utilize ‘Orah’ mandarin as the female parent.

• Data here indicate that ‘Orah’ mandarin is a cultivar that readily produces 2n megagametophytes.

• Second division restitution is the mechanism underlying 2n megagametophyte formation in ‘Orah’ mandarin.

Seedless fruits are desirable in the citrus fresh fruit market. Triploid production via diploid × tetraploid interploidy crosses is thought to be the most efficient and widely-used strategy for the breeding of seedless citrus. Although ‘Orah’ mandarin has desirable organoleptic qualities, seeds in the fruits weaken its market competitiveness. To produce new seedless cultivars that are similar to ‘Orah’ mandarin, we performed three 2x × 4x crosses using ‘Orah’ mandarin as the seed parent to regenerate triploid plantlets. A total of 182 triploid and 36 tetraploid plantlets were obtained. By analyzing their genetic origins using nine novel single nucleotide polymorphism (SNP) markers, all of the triploids and tetraploids derived from these three crosses were proven to be hybrids. Also, we demonstrated that 2n megagametophyte formation in ‘Orah’ mandarin result in tetraploid production in these three interploidy crosses. These tetraploid plantlets were genotyped using eight pericentromeric SNP markers and nine centromere distal SNP markers. Based on the genotypes of the 2n megagametophytes, the parental heterozygosity rates in 16 SNP loci and all 2n megagametophytes were less than 50%, indicating that second division restitution was the mechanism underlying 2n megagametophyte formation at both the population and individual levels. These triploid hybrids enrich the germplasm available for seedless breeding. Moreover, the tetraploid hybrids are valuable as parents for ploidy breeding for the production of seedless citrus fruits.
Keywords Citrus      2n gamete      interploidy hybridization      pericentromeric SNP marker      second division restitution     
Corresponding Author(s): Kaidong XIE,Wenwu GUO   
Just Accepted Date: 03 February 2021   Online First Date: 15 March 2021    Issue Date: 13 July 2021
 Cite this article:   
Qiangming XIA,Wei WANG,Kaidong XIE, et al. UNREDUCED MEGAGAMETOPHYTE FORMATION VIA SECOND DIVISION RESTITUTION CONTRIBUTES TO TETRAPLOID PRODUCTION IN INTERPLOIDY CROSSES WITH ‘ORAH’ MANDARIN (CITRUS RETICULATA)[J]. Front. Agr. Sci. Eng. , 2021, 8(2): 302-313.
 URL:  
https://academic.hep.com.cn/fase/EN/10.15302/J-FASE-2021385
https://academic.hep.com.cn/fase/EN/Y2021/V8/I2/302
Cross No. pollinated flowers No. fruits set No. seeds obtained No. seeds germinated No. plantlets obtained No. diploids No. triploids No. tetraploids
Dev. Undev. Dev. Undev.
Orah × PCS 210 79 323 930 115 271 145 37 90 18
Orah × PO 238 128 859 906 99 108 132 35 81 16
Orah × SP 263 80 490 511 101 87 88 75 11 2
Total 711 287 1672 2347 315 466 365 147 182 36
Tab.1  The fruit set and numbers of seeds and polyploids recovered from the 2x × 4x crosses
Fig.1  Embryo rescue, plant regeneration and transplantation for citrus triploid production. (a) Young fruits 85 d after pollination. (b) Germination of developed seeds after approximately two weeks of culturing in vitro on germination medium. (c) Germination of undeveloped seeds after about four weeks of culturing in vitro on germination medium. (d) Regeneration of shoots from embryoids after their transfer to the shoot-induction medium. (e) A shoot grafted in vitro to the rootstock (Poncirus trifoliata). (f) Transplanted seedlings in a greenhouse.
Fig.2  Ploidy determination for regenerated citrus plantlets using flow cytometry and chromosome counting. (a–c) Histograms of diploid progeny (peak= 50), triploid progeny (peak= 75) and tetraploid progeny (peak= 100). (d–f) Chromosome counting for diploid (2n= 2x= 18), triploid (2n= 3x= 27) and tetraploid (2n= 4x= 36) plantlets. Scale bars= 5 mm.
Fig.3  Determining the genetic origin of triploids and tetraploids using KASP genotyping and aa × bbbb type SNP markers. Genotyping plots of (a) 43 randomly selected triploid progeny and (b) 36 tetraploid progeny with SNP marker Chr2-25841537 demonstrating their hybrid origins. Green, blue, red and gray represent the genotypes of maternal parents, paternal parents, triploid or tetraploid progeny and negative controls, respectively.
SNP marker Orah (aa) Male parents (bbbb) NI abb aabb
Chr2-24850985 CC AAAA 79 43 36
Chr2-25841537 GG CCCC 79 43 36
Chr3-18395328 AA GGGG 78 43 35
Chr3-24832283 CC TTTT 79 43 36
Chr4-8664085 GG CCCC 77 42 35
Chr4-8689111 GG TTTT 79 43 36
Chr5-12876197 CC TTTT 79 43 36
Chr6-1932038 TT CCCC 79 43 36
Chr9-815315 AA GGGG 76 43 33
Tab.2  Genotypic analysis of nine SNP markers (aa × bbbb type) in the triploid and tetraploid hybrid populations
Fig.4  Determining the mechanism of 2n megagametophyte formation in the 36 tetraploids using KASP genotyping and ab × aaaa/bbbb type SNP markers. (a) Under pericenteomeric locus Chr5-17395118, the maternal genotype (green) is GA, the paternal genotype (blue) is GGGG, and the tetraploid plantlets (red) clustered with their parents; the genotypes of the tetraploids are GGAA and GGGG with a GG contribution from the paternal parent and therefore homozygous AA and GG for the 2n megagametophyte. (b) Under the centromere distal locus Chr5-24798525, the maternal genotype (green) is TC, the paternal genotype (blue) is TTTT, and the tetraploid plantlets (red) clustered into three groups; the genotypes of the tetraploids are TTCC, TTTT and TTTC with a TT contribution from the paternal parent and therefore homozygous CC, TT and TC for the 2n megagametophyte.
SNP
markers		 Orah Male parents OPCS
1		 OPCS
2		 OPCS
3		 OPCS
4		 OPCS
5		 OPCS
6		 OPCS
7		 OPCS
8		 OPCS
9		 OPCS
10		 OPCS
11		 OPCS
12		 OPCS
13		 OPCS
14		 OPCS
15		 OPCS
16		 OPCS
17		 OPCS
18		 Het PHR
Chr1-12169985 GA GGGG GG AA GG GG AA AA GG GG GG AA AA GG GG AA GG AA AA GG 0 0
Chr3-7913461 GA GGGG GG AA GG GG AA AA GG AA AA AA AA GG AA GG AA GG GG GG 0 0
Chr5-17395118 GA GGGG GG AA AA GG AA GG AA AA GG AA AA GG AA GG AA GG GG AA 0 0
Chr5-18735709 TA TTTT TT TT AA TT TT AA TT TT TT TT TT AA TT AA TT TT TT TT 0 0
Chr6-5337355 TG TTTT TT TT GG GG GG TT GG TT TT TT GG GG TT TT GG GG GG TT 0 0
Chr7-20190172 CA CCCC CC AA CC AA AA CC AA AA CC CC CC CC CC AA CC AA AA AA 0 0
Chr8-7202641 AG AAAA AA AA GG AA GG AA AA AA GG GG AA GG AA AA GG GG GG GG 0 0
Chr9-8606082 GT GGGG TT GG GG GG TT TT GG GG GG TT GG TT TT GG TT GG TT GG 0 0
Chr5-1014992 CA CCCC AA CC AA AA CC AA AA AA AA AA AA AA AA CC AA CC AA AA 0 0
Chr5-1051787 AT AAAA TT AA AA AA AA TT TT TT AA TT TT AA AA AA AA AA AA AA 0 0
Chr5-1103777 TC TTTT TC TC TC TC TT TC TC TC TC TC TC CC TC TT TC TC TC TC 15 83.33
Chr5-1323430 CT CCCC TT CC CC CC CC TT TT TT TT TT TT CC CC CC CC CC CC TT 0 0
Chr5-1580076 TC TTTT CC TT CC CC TT CC CC CC CC CC CC CC CC TT CC TT CC TT 0 0
Chr5-22348846 GC GGGG GG GG GG CC CC GG CC CC CC CC GG GG GG CC GG CC GG CC 0 0
Chr5-24661722 AG AAAA GG AA GG GG GG GG GG GG AA GG GG GG GG AA GG AA GG AA 0 0
Chr5-24798525 TC TTTT TT TT TT TC TC TC TC CC TT TT TC TC TT TC TC CC TT TT 8 44.44
Chr5-26120337 CT CCCC CC CC TT TT CC TT TT TT TT CC TT TT TT CC TT CC TT CC 0 0
Het 1 1 1 2 1 2 2 1 1 1 2 1 1 1 2 1 1 1
PHR 5.88 5.88 5.88 11.76 5.88 11.76 11.76 5.88 5.88 5.88 11.76 5.88 5.88 5.88 11.76 5.88 5.88 5.88
Tab.3  Genotypes of 18 tetraploids from ‘Orah × PCS’ hybridization generated using eight pericentromeric SNP markers and nine centromere distal SNP markers
SNP markers Orah Male parents OPO1 OPO2 OPO3 OPO4 OPO5 OPO6 OPO7 OPO8 OPO9 OPO10 OPO11 OPO12 OPO13 OPO14 OPO15 OPO16 OSP1 OSP2 Het PHR
Chr1-12169985 GA GGGG GG GG AA GG AA AA AA AA GG GG AA AA GG GG AA GG AA AA 0 0
Chr3-7913461 GA GGGG AA AA GG AA GG GG GG AA GG GG GG AA AA GG GG AA GG GG 0 0
Chr5-17395118 GA GGGG AA AA GG AA GG AA GG AA AA GG GG AA GG AA AA GG AA AA 0 0
Chr5-18735709 TA TTTT AA AA TT AA TT AA AA AA TT TT TT AA TT AA TT TT TT TT 0 0
Chr6-5337355 TG TTTT TT TT TT GG TT GG GG TT TT GG TT GG GG GG TT GG GG GG 0 0
Chr7-20190172 CA CCCC CC CC CC AA CC CC AA CC CC AA CC CC CC AA AA AA CC CC 0 0
Chr8-7202641 AG AAAA AA AA AA AA GG AA GG AA AA AA AA GG GG AA AA GG GG GG 0 0
Chr9-8606082 GT GGGG GG GG GG TT TT GG GG GG TT TT TT GG GG GG TT GG GG GG 0 0
Chr5-1014992 CA CCCC CC AA AA CC CC CC CC CC CC AA AA CC AA CC AA CC CC AA 0 0
Chr5-1051787 AT AAAA AA AA AA AA AA AA AA AA TT AA AA AA AA AA AA AA AA AA 0 0
Chr5-1103777 TC TTTT TC TC TC TC TC TC TC TC TC TC TC TC TC TC TC TT TT TC 16 88.89
Chr5-1323430 CT CCCC CC CC CC CC CC CC CC CC CC TT CC CC CC CC CC CC CC CC 0 0
Chr5-1580076 TC TTTT TT CC CC TT TT TT TT TT TT CC CC TT CC TT CC TT TT CC 0 0
Chr5-22348846 GC GGGG GG CC CC GG GG GG GG GG GG CC GG GG GG GG CC CC CC GG 0 0
Chr5-24661722 AG AAAA AA GG AA AA AA AA AA AA AA AA GG AA GG AA GG GG GG GG 0 0
Chr5-24798525 TC TTTT TT TC CC TT TT TT TT TT TT CC TT TT TT TT TC TC TC TC 5 27.78
Chr5-26120337 CT CCCC CC TT CC CC CC CC CC CC CC TT TT CC TT CC CC TT CC CC 0 0
Het 1 2 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 2
PHR 5.88 11.76 5.88 5.88 5.88 5.88 5.88 5.88 5.88 5.88 5.88 5.88 5.88 5.88 11.76 5.88 5.88 11.76
Tab.4  Genotypes of 18 tetraploids from ‘Orah × PO’ and ‘Orah × SP’ hybridizations generated using eight pericentromeric SNP markers and nine centromere distal SNP markers
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