1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.
21.
22.
23.
24.
25.
26.
27.
28.
29.
30.
31.
Darwin, C. The Effects of Cross and Self Fertilisation in the Vegetable Kingdom, 1st ed.; John Murray: London, UK, 1876; pp. 1–482.
Shull, G.H. What is “Heterosis”? Genetics 1948, 33, 439–446. [CrossRef] [PubMed]
Crow, J.F. 90 years ago: The beginning of hybrid maize. Genetics 1998, 148, 923–928. [CrossRef] [PubMed]
Ma, G.H.; Yuan, L.P. Hybrid rice achievements, development and prospect in China. J. Integr. Agric. 2015, 14, 197–205. [CrossRef]
Wakchaure, R.; Ganguly, S.; Praveen, P.K.; Sharma, S.; Kumar, A.; Mahajan, T.; Qadri, K. Importance of heterosis in animals: A
review. Int. J. Adv. Eng. Technol. Innov. Sci. 2015, 1, 1–5.
Bruce, A.B. The mendelian theory of heredity and the augmentation of vigor. Science 1910, 32, 627–628. [CrossRef]
Jones, D.F. Dominance of linked factors as a means of accounting for heterosis. Proc. Natl. Acad. Sci. USA 1917, 3, 310–312.
[CrossRef]
Hashimoto, S.; Wake, T.; Nakamura, H.; Minamiyama, M.; Araki-Nakamura, S.; Ohmae-Shinohara, K.; Koketsu, E.; Okamura, S.;
Miura, K.; Kawaguchi, H.; et al. The dominance model for heterosis explains culm length genetics in a hybrid sorghum variety.
Sci. Rep. 2021, 11, 4532. [CrossRef]
Busch, R.H.; Luchen, K.A.; Frohberg, R.C. F1 hybrids versus random F5 line performance and estimates of genetic effects in
spring wheat. Crop Sci. 1971, 11, 357–361. [CrossRef]
Wang, L.; Greaves, I.K.; Groszmann, M.; Wu, L.M.; Dennis, E.S.; Peacock, W.J. Hybrid mimics and hybrid vigor in Arabidopsis.
Proc. Natl. Acad. Sci. USA 2015, 112, E4959–E4967. [CrossRef]
He, Y.; Zhang, Y.; Liao, Y.; Dennis, E.S.; Peacock, W.J.; Wu, X. Rice hybrid mimics have stable yields equivalent to those of the F1
hybrid and suggest a basis for hybrid vigour. Planta 2021, 254, 51. [CrossRef]
Zhang, Y.; Ovenden, B.; He, Y.; Ye, W.; Wu, X.; Peacock, W.J.; Dennis, E.S. Hybrid vigour and hybrid mimics in Japonica rice.
Agronomy 2022, 12, 1559. [CrossRef]
Hull, F.H. Recurrent selection and overdominance. In Heterosis; Gowen, J.W., Ed.; Iowa State College Press: Ames, IA, USA, 1952;
pp. 451–473.
Krieger, U.; Lippman, Z.B.; Zamir, D. The flowering gene SINGLE FLOWER TRUSS drives heterosis for yield in tomato. Nat.
Genet. 2010, 42, 459–463. [CrossRef]
Charlesworth, D.; Willis, J. The genetics of inbreeding depression. Nat. Rev. Genet. 2009, 10, 783–796. [CrossRef]
Yu, D.; Gu, X.; Zhang, S.; Dong, S.; Miao, H.; Gebretsadik, K.; Bo, K. Molecular basis of heterosis and related breeding strategies
reveal its importance in vegetable breeding. Hortic. Res. 2021, 8, 120. [CrossRef]
Guo, T.; Yang, N.; Tong, H.; Pan, Q.; Yang, X.; Tang, J.; Wang, J.; Li, J.; Yan, J. Genetic basis of grain yield heterosis in an
“immortalized F2 ” maize population. Theor. Appl. Genet. 2014, 127, 2149–2158. [CrossRef]
Huang, X.; Yang, S.; Gong, J.; Zhao, Q.; Feng, Q.; Zhan, Q.; Zhao, Y.; Li, W.; Cheng, B.; Xia, J.; et al. Genomic architecture of
heterosis for yield traits in rice. Nature 2016, 537, 629–633. [CrossRef]
Fujimoto, R.; Uezono, K.; Ishikura, S.; Osabe, K.; Peacock, W.J.; Dennis, E.S. Recent research on the mechanism of heterosis is
important for crop and vegetable breeding systems. Breed. Sci. 2018, 68, 145–158. [CrossRef]
Fortuny, A.P.; Bueno, R.A.; Pereira da Costa, J.H.; Zanor, M.I.; Rodríguez, G.R. Tomato fruit quality traits and metabolite content
are affected by reciprocal crosses and heterosis. J. Exp. Bot. 2021, 72, 5407–5425. [CrossRef]
Kaushik, P.; Plazas, M.; Prohens, J.; Vilanova, S.; Gramazio, P. Diallel genetic analysis for multiple traits in eggplant and assessment
of genetic distances for predicting hybrids performance. PLoS ONE 2018, 13, e0199943. [CrossRef]
Yue, L.; Zhang, S.; Zhang, L.; Liu, Y.; Cheng, F.; Li, G.; Zhang, S.; Zhang, H.; Sun, R.; Li, F. Heterotic prediction of hybrid
performance based on genome-wide SNP markers and the phenotype of parental inbred lines in heading Chinese cabbage
(Brassica rapa L. ssp. pekinensis). Sci. Hortic. 2022, 296, 110907. [CrossRef]
Kawamura, K.; Kawanabe, T.; Shimizu, M.; Nagano, A.J.; Saeki, N.; Okazaki, K.; Kaji, M.; Dennis, E.S.; Osabe, K.; Fujimoto, R.
Genetic distance of inbred lines of Chinese cabbage and its relationship to heterosis. Plant Gene 2016, 5, 1–7. [CrossRef]
Chen, Z.J. Genomic and epigenetic insights into the molecular bases of heterosis. Nat. Rev. Genet. 2013, 14, 471–482. [CrossRef]
[PubMed]
Lippman, Z.B.; Zamir, D. Heterosis: Revisiting the magic. Trends Genet. 2007, 23, 60–66. [CrossRef]
Groszmann, M.; Greaves, I.K.; Fujimoto, R.; Peacock, W.J.; Dennis, E.S. The role of epigenetics in hybrid vigour. Trends Genet.
2013, 29, 684–690. [CrossRef] [PubMed]
Santamaria, P.; Signore, A. How has the consistency of the common catalogue of varieties of vegetable species changed in the last
ten years? Sci. Hortic. 2021, 277, 109805. [CrossRef]
Yamagishi, H.; Bhat, S.R. Cytoplasmic male sterility in Brassicaceae crops. Breed. Sci. 2014, 64, 38–47. [CrossRef]
Fujimoto, R.; Nishio, T. Self-Incompatibility. Adv. Bot. Res. 2007, 45, 139–154.
Chen, L.; Liu, Y.G. Male sterility and fertility restoration in crops. Annu. Rev. Plant Biol. 2014, 65, 579–606. [CrossRef]
Singh, S.; Dey, S.S.; Bhatia, R.; Kumar, R.; Behera, T.K. Current understanding of male sterility systems in vegetable Brassicas and
their exploitation in hybrid breeding. Plant Reprod. 2019, 32, 231–256. [CrossRef]
Horticulturae 2023, 9, 366
32.
33.
34.
35.
36.
37.
38.
39.
40.
41.
42.
43.
44.
45.
46.
47.
48.
49.
50.
51.
52.
53.
54.
55.
56.
14 of 18
Seymour, D.K.; Chae, E.; Grimm, D.G.; Pizarro, C.M.; Habring-Müller, A.; Vasseur, F.; Rakitsch, B.; Borgwardt, K.M.; Koenig, D.;
Weigel, D. Genetic architecture of nonadditive inheritance in Arabidopsis thaliana hybrids. Proc. Natl. Acad. Sci. USA 2016, 113,
E7317–E7326. [CrossRef]
Yang, M.; Wang, X.; Ren, D.; Huang, H.; Xu, M.; He, G.; Deng, X.W. Genomic architecture of biomass heterosis in Arabidopsis.
Proc. Natl. Acad. Sci. USA 2017, 114, 8101–8106. [CrossRef]
Kusterer, B.; Piepho, H.P.; Utz, H.F.; Schön, C.C.; Muminovic, J.; Meyer, R.C.; Altmann, T.; Melchinger, A.E. Heterosis for
biomass-related traits in Arabidopsis investigated by quantitative trait loci analysis of the triple testcross design with recombinant
inbred lines. Genetics 2007, 177, 1839–1850. [CrossRef]
Meyer, R.C.; Kusterer, B.; Lisec, J.; Steinfath, M.; Becher, M.; Scharr, H.; Melchinger, A.E.; Selbig, J.; Schurr, U.; Willmitzer, L.; et al.
QTL analysis of early stage heterosis for biomass in Arabidopsis. Theor. Appl. Genet. 2010, 120, 227–237. [CrossRef]
Lisec, J.; Steinfath, M.; Meyer, R.C.; Selbig, J.; Melchinger, A.E.; Willmitzer, L.; Altmann, T. Identification of heterotic metabolite
QTL in Arabidopsis thaliana RIL and IL populations. Plant J. 2009, 59, 777–788. [CrossRef]
Andorf, S.; Meyer, R.C.; Selbig, J.; Altmann, T.; Repsilber, D. Integration of a systems biological network analysis and QTL results
for biomass heterosis in Arabidopsis thaliana. PLoS ONE 2012, 7, e49951. [CrossRef]
Fujimoto, R.; Taylor, J.M.; Shirasawa, S.; Peacock, W.J.; Dennis, E.D. Heterosis of Arabidopsis hybrids between C24 and Col is
associated with increased photosynthesis capacity. Proc. Natl. Acad. Sci. USA 2012, 109, 7109–7114. [CrossRef]
Saeki, N.; Kawanabe, T.; Ying, H.; Shimizu, M.; Kojima, M.; Abe, H.; Okazaki, K.; Kaji, M.; Taylor, J.M.; Sakakibara, H.; et al.
Molecular and cellular characteristics of hybrid vigour in a commercial hybrid of Chinese cabbage. BMC Plant Biol. 2016, 16, 45.
[CrossRef]
Ni, Z.; Kim, E.D.; Ha, M.; Lackey, E.; Liu, J.; Zhang, Y.; Sun, Q.; Chen, Z.J. Altered circadian rhythms regulate growth vigour in
hybrids and allopolyploids. Nature 2009, 457, 327–331. [CrossRef]
Shen, H.; He, H.; Li, J.; Chen, W.; Wang, X.; Guo, L.; Peng, Z.; He, G.; Zhong, S.; Qi, Y.; et al. Genome-wide analysis of DNA
methylation and gene expression changes in two Arabidopsis ecotypes and their reciprocal hybrids. Plant Cell 2012, 24, 875–892.
[CrossRef]
Groszmann, M.; Gonzalez-Bayon, R.; Lyons, R.L.; Greaves, I.K.; Kazan, K.; Peacock, W.J.; Dennis, E.S. Hormone-regulated defense
and stress response networks contribute to heterosis in Arabidopsis F1 hybrids. Proc. Natl. Acad. Sci. USA 2015, 112, E6397–E6406.
[CrossRef]
Alonso-Peral, M.M.; Trigueros, M.; Sherman, B.; Ying, H.; Taylor, J.M.; Peacock, W.J.; Dennis, E.S. Patterns of gene expression in
developing embryos of Arabidopsis hybrids. Plant J. 2017, 89, 927–939. [CrossRef] [PubMed]
Wang, L.; Wu, L.M.; Greaves, I.K.; Zhu, A.; Dennis, E.S.; Peacock, W.J. PIF4-controlled auxin pathway contributes to hybrid vigor
in Arabidopsis thaliana. Proc. Natl. Acad. Sci. USA 2017, 114, E3555–E3562. [CrossRef] [PubMed]
Gonzalez-Bayon, R.; Shen, Y.; Groszmann, M.; Zhu, A.; Wang, A.; Allu, A.D.; Dennis, E.S.; Peacock, W.J.; Greaves, I.K. Senescence
and defense pathways contribute to heterosis. Plant Physiol. 2019, 180, 240–252. [CrossRef] [PubMed]
Zhu, A.; Greaves, I.K.; Liu, P.C.; Wu, L.; Dennis, E.S.; Peacock, W.J. Early changes of gene activity in developing seedlings of
Arabidopsis hybrids relative to parents may contribute to hybrid vigour. Plant J. 2016, 88, 597–607. [CrossRef]
Meyer, R.C.; Witucka-Wall, H.; Becher, M.; Blacha, A.; Boudichevskaia, A.; Dörmann, P.; Fiehn, O.; Friedel, S.; von Korff, M.; Lisec,
J.; et al. Heterosis manifestation during early Arabidopsis seedling development is characterized by intermediate gene expression
and enhanced metabolic activity in the hybrids. Plant J. 2012, 71, 669–683. [CrossRef]
Liu, W.; He, G.; Deng, X.W. Biological pathway expression complementation contributes to biomass heterosis in Arabidopsis.
Proc. Natl. Acad. Sci. USA 2021, 118, e2023278118. [CrossRef]
Dominissini, D.; Moshitch-Moshkovitz, S.; Schwartz, S.; Salmon-Divon, M.; Ungar, L.; Osenberg, S.; Cesarkas, K.; Jacob-Hirasch,
J.; Amariglio, N.; Kupiec, M.; et al. Topology of the human and mouse m6 A RNA methylomes revealed by m6 A-seq. Nature 2012,
485, 201–206. [CrossRef]
Meyer, K.D.; Saletore, Y.; Zumbo, P.; Elemento, O.; Mason, C.E.; Jaffrey, S.R. Comprehensive analysis of mRNA methylation
reveals enrichment in 3’ UTRs and near stop codons. Cell 2012, 149, 1635–1646. [CrossRef]
Yue, H.; Nie, X.; Yan, Z.; Weining, S. N6-methyladenosine regulatory machinery in plants: Composition, function and evolution.
Plant Biotechnol. J. 2019, 17, 1194–1208. [CrossRef]
Xu, Z.; Shi, X.; Bao, M.; Song, X.; Zhang, Y.; Wang, H.; Xie, H.; Mao, F.; Wang, S.; Jin, H.; et al. Transcriptome-wide analysis of
RNA m6 A methylation and gene expression changes among two Arabidopsis ecotypes and their reciprocal hybrids. Front. Plant
Sci. 2021, 12, 685189. [CrossRef]
Fujimoto, R.; Sasaki, T.; Ishikawa, R.; Osabe, K.; Kawanabe, T.; Dennis, E.S. Molecular mechanisms of epigenetic variation in
plants. Int. J. Mol. Sci. 2012, 13, 9900–9922. [CrossRef]
Kawakatsu, T.; Ecker, J.R. Diversity and dynamics of DNA methylation: Epigenomic resources and tools for crop breeding. Breed.
Sci. 2019, 69, 191–204. [CrossRef]
Greaves, I.K.; Groszmann, M.; Ying, H.; Taylor, J.M.; Peacock, W.J.; Dennis, E.S. Trans chromosomal methylation in Arabidopsis
hybrids. Proc. Natl. Acad. Sci. USA 2012, 109, 3570–3575. [CrossRef]
Zhu, A.; Greaves, I.K.; Dennis, E.S.; Peacock, W.J. Genome-wide analyses of four major histone modifications in Arabidopsis
hybrids at the germinating seed stage. BMC Genom. 2017, 18, 137. [CrossRef]
Horticulturae 2023, 9, 366
57.
58.
59.
60.
61.
62.
63.
64.
65.
66.
67.
68.
69.
70.
71.
72.
73.
74.
75.
76.
77.
78.
79.
80.
81.
82.
83.
15 of 18
Zhang, Q.; Wang, D.; Lang, Z.; He, L.; Yang, L.; Zeng, L.; Li, Y.; Zhao, C.; Huang, H.; Zhang, H.; et al. Methylation interactions in
Arabidopsis hybrids require RNA-directed DNA methylation and are influenced by genetic variation. Proc. Natl. Acad. Sci. USA
2016, 113, E4248–E4256. [CrossRef]
Rigal, M.; Becker, C.; Pélissier, T.; Pogorelcnik, R.; Devos, J.; Ikeda, Y.; Weigel, D.; Mathieu, O. Epigenome confrontation triggers
immediate reprogramming of DNA methylation and transposon silencing in Arabidopsis thaliana F1 epihybrids. Proc. Natl. Acad.
Sci. USA 2016, 113, E2083–E2092. [CrossRef]
Kawanabe, T.; Ishikura, S.; Miyaji, N.; Sasaki, T.; Wu, L.M.; Itabashi, E.; Takada, S.; Shimizu, M.; Takasaki-Yasuda, T.; Osabe,
K.; et al. Role of DNA methylation in hybrid vigor in Arabidopsis thaliana. Proc. Natl. Acad. Sci. USA 2016, 113, E6704–E6711.
[CrossRef]
Dapp, M.; Reinders, J.; Bédiée, A.; Balsera, C.; Bucher, E.; Theiler, G.; Granier, C.; Paszkowski, J. Heterosis and inbreeding
depression of epigenetic Arabidopsis hybrids. Nat. Plants 2015, 1, 15092. [CrossRef]
Lauss, K.; Wardenaar, R.; Oka, R.; van Hulten, M.H.A.; Guryev, V.; Keurentjes, J.J.B.; Stam, M.; Johannes, F. Parental DNA
methylation states are associated with heterosis in epigenetic hybrids. Plant Physiol. 2018, 176, 1627–1645. [CrossRef]
Zhang, Q.; Li, Y.; Xu, T.; Srivastava, A.K.; Wang, D.; Zeng, L.; Yang, L.; He, L.; Zhang, H.; Zheng, Z.; et al. The chromatin
remodeler DDM1 promotes hybrid vigor by regulating salicylic acid metabolism. Cell Discov. 2016, 2, 16027. [CrossRef]
Miyaji, N.; Fujimoto, R. Hybrid vigor: Importance of epigenetic processes and consequences for breeding. Adv. Bot. Res. 2018, 88,
247–275.
Miller, M.; Song, Q.; Shi, X.; Juenger, E.T.; Chen, Z.J. Natural variation in timing of stress-responsive gene expression predicts
heterosis in intraspecific hybrids of Arabidopsis. Nat. Commun. 2015, 6, 7453. [CrossRef] [PubMed]
Yang, L.; Li, B.; Zheng, X.Y.; Li, J.; Yang, M.; Dong, X.; He, G.; An, C.; Deng, X.W. Salicylic acid biosynthesis is enhanced and
contributes to increased biotrophic pathogen resistance in Arabidopsis hybrids. Nat. Commun. 2015, 6, 7309.
Calvo-Baltanás, V.; Wang, J.; Chae, E. Hybrid incompatibility of the plant immune system: An opposite force to heterosis
equilibrating hybrid performances. Front. Plant Sci. 2021, 11, 576796. [CrossRef]
Yang, L.; Liu, P.; Wang, X.; Jia, A.; Ren, D.; Tang, Y.; Tang, Y.; Deng, X.W.; He, G. A central circadian oscillator confers defense
heterosis in hybrids without growth vigor costs. Nat. Commun. 2021, 12, 2317. [CrossRef]
Shull, G.H. The composition of a field of maize. J. Hered. 1908, 4, 296–301. [CrossRef]
East, E.M.; Jones, D.F. Inbreeding and Outbreeding: Their Genetic and Sociological Significance; Lippincott: Philadelphia, PA, USA,
1919; p. 285.
Hochholdinger, F.; Hoecker, N. Towards the molecular basis of heterosis. Trends Plant Sci. 2007, 12, 427–432. [CrossRef]
Barth, S.; Busimi, A.K.; Friedrich Utz, H.; Melchinger, A.E. Heterosis for biomass yield and related traits in five hybrids of
Arabidopsis thaliana L. Heynh. Heredity 2003, 91, 36–42. [CrossRef]
Meyer, R.C.; Törjék, O.; Becher, M.; Altmann, T. Heterosis of biomass production in Arabidopsis. Establishment during early
development. Plant Physiol. 2004, 134, 1813–1823. [CrossRef]
Syed, N.H.; Chen, Z.J. Molecular marker genotypes, heterozygosity and genetic interactions explain heterosis in Arabidopsis
thaliana. Heredity 2005, 94, 295–304. [CrossRef]
Groszmann, M.; Gonzalez-Bayon, R.; Greaves, I.K.; Wang, L.; Huen, A.K.; Peacock, W.J.; Dennis, E.S. Intraspecific Arabidopsis
hybrids show different patterns of heterosis despite the close relatedness of the parental genomes. Plant Physiol. 2014, 166,
265–280. [CrossRef]
van Hulten, M.H.A.; Paulo, M.J.; Kruijer, W.; Vries, H.B.D.; Kemperman, B.; Becker, F.F.M.; Yang, J.; Lauss, K.; Stam, M.E.; van
Eeuwijk, F.A.; et al. Assessment of heterosis in two Arabidopsis thaliana common-reference mapping populations. PLoS ONE 2018,
13, e0205564. [CrossRef]
Wang, L.; Wu, L.M.; Greaves, I.K.; Dennis, E.S.; Peacock, W.J. In Arabidopsis hybrids and Hybrid Mimics, up-regulation of cell
wall biogenesis is associated with the increased plant size. Plant Direct 2019, 3, e00174. [CrossRef]
Li, P.; Su, T.; Zhang, D.; Wang, W.; Xin, X.; Yu, Y.; Zhao, X.; Yu, S.; Zhang, F. Genome-wide analysis of changes in miRNA and
target gene expression reveals key roles in heterosis for Chinese cabbage biomass. Hortic. Res. 2021, 8, 39. [CrossRef]
Jeong, S.Y.; Ahmed, N.U.; Jung, H.J.; Kim, H.T.; Park, J.I.; Nou, I.S. Discovery of candidate genes for heterosis breeding in Brassica
oleracea L. Acta Physiol. Plant. 2017, 39, 180. [CrossRef]
Verma, V.K.; Kalia, P. Combining ability analysis and its relationship with gene action and heterosis in early maturity cauliflower.
Proc. Natl. Acad. Sci. India Sect. B Biol. Sci. 2017, 87, 877–884. [CrossRef]
Basunanda, P.; Radoev, M.; Ecke, W.; Friedt, W.; Becker, H.C.; Snowdon, R.J. Comparative mapping of quantitative trait loci
involved in heterosis for seedling and yield traits in oilseed rape (Brassica napus L.). Theor. Appl. Genet. 2010, 120, 271–280.
[CrossRef]
Wolko, J.; Dobrzycka, A.; Bocianowski, J.; Bartkowiak-Broda, I. Estimation of heterosis for yield-related traits for single cross and
three-way cross hybrids of oilseed rape (Brassica napus L.). Euphytica 2019, 215, 156. [CrossRef]
Zhu, A.; Wang, A.; Zhang, Y.; Dennis, E.S.; Peacock, W.J.; Greaves, A. Early establishment of photosynthesis and auxin biosynthesis
plays a key role in early biomass heterosis in Brassica napus (Canola) hybrids. Plant Cell Physiol. 2020, 61, 1134–1143. [CrossRef]
Aakanksha; Yadava, S.K.; Yadav, B.G.; Gupta, V.; Mukhopadhyay, A.; Pental, D.; Pradhan, A.K. Genetic analysis of heterosis for
yield influencing traits in Brassica juncea using a doubled haploid population and its backcross progenies. Front. Plant Sci. 2021,
12, 721631. [CrossRef]
Horticulturae 2023, 9, 366
84.
85.
86.
87.
88.
89.
90.
91.
92.
93.
94.
95.
96.
97.
98.
99.
100.
101.
102.
103.
104.
105.
106.
107.
108.
109.
16 of 18
Wang, Q.; Yan, T.; Long, Z.; Huang, L.Y.; Zhu, Y.; Xu, Y.; Chen, X.; Pak, H.; Li, J.; Wu, D.; et al. Prediction of heterosis in the
recent rapeseed (Brassica napus) polyploid by pairing parental nucleotide sequences. PLoS Genet. 2021, 17, e1009879. [CrossRef]
[PubMed]
Hu, Y.; Xiong, J.; Shalby, N.; Zhuo, C.; Jia, Y.; Yang, Q.Y.; Tu, J. Comparison of dynamic 3D chromatin architecture uncovers
heterosis for leaf size in Brassica napus. J. Adv. Res. 2022, 42, 289–301. [CrossRef] [PubMed]
Tamta, S.; Singh, J. Heterosis in tomato for growth and yield traits. Int. J. Veg. Sci. 2018, 24, 169–179. [CrossRef]
Chandel, R.; Sadashiva, A.T.; Ravishankar, K.V.; Das, A.; Rout, B.M.; Singh, S. Genetic combining, heterosis analysis for
horticultural traits in tomato (Solanum lycopersicum L.) using ToLCV-resistant lines and molecular validation of Ty genes. Plant
Genet. Resour. 2021, 19, 512–521. [CrossRef]
Rajendran, S.; Bae, J.H.; Park, M.W.; Oh, J.H.; Jeong, H.W.; Lee, Y.K.; Park, S.J. Tomato yield effects of reciprocal hybridization of
Solanum lycopersicum cultivars M82 and Micro-Tom. Plant Breed. Biotechnol. 2022, 10, 37–48. [CrossRef]
Kakizaki, Y. Hybrid vigor in egg-plants and its practical utilization. Genetics 1931, 16, 1–25. [CrossRef]
Kaushik, P.; Prohens, J.; Vilanova, S.; Gramazio, P.; Plazas, M. Phenotyping of eggplant wild relatives and interspecific hybrids
with conventional and phenomics descriptors provides insight for their potential utilization in breeding. Front. Plant Sci. 2016, 7,
677. [CrossRef]
Kaushik, P. Line × Tester analysis for morphological and fruit biochemical traits in eggplant (Solanum melongena L.) using wild
relatives as testers. Agronomy 2019, 9, 185. [CrossRef]
Kumar, A.; Sharma, V.; Jain, B.T.; Kaushik, P. Heterosis breeding in eggplant (Solanum melongena L.): Gains and provocations.
Plants 2020, 9, 403. [CrossRef]
Tian, F.; Bradbury, P.J.; Brown, P.J.; Hung, H.; Sun, Q.; Flint-Garcia, S.; Rocheford, T.R.; McMullen, M.D.; Holland, J.B.; Buckler,
E.S. Genome-wide association study of leaf architecture in the maize nested association mapping population. Nat. Genet. 2011, 43,
159–162. [CrossRef]
Ko, D.K.; Rohozinski, D.; Song, Q.; Taylor, S.H.; Juenger, T.E.; Harmon, F.G.; Chen, Z.J. Temporal shift of circadian-mediated gene
expression and carbon fixation contributes to biomass heterosis in maize hybrids. PLoS Genet. 2016, 12, e1006197. [CrossRef]
Song, Y.; Zhang, Z.; Tan, X.; Jiang, Y.; Gao, J.; Lin, L.; Wang, Z.; Ren, J.; Wang, X.; Qin, L.; et al. Association of the molecular
regulation of ear leaf senescence/stress response and photosynthesis/metabolism with heterosis at the reproductive stage in
maize. Sci. Rep. 2016, 6, 29843. [CrossRef]
Yang, J.; Mezmouk, S.; Baumgarten, A.; Buckler, E.S.; Guill, K.E.; McMullen, M.D.; Mumm, R.H.; Ross-Ibarra, J. Incomplete
dominance of deleterious alleles contributes substantially to trait variation and heterosis in maize. PLoS Genet. 2017, 13, e1007019.
[CrossRef]
Liu, H.; Wang, Q.; Chen, M.; Ding, Y.; Yang, X.; Liu, J.; Li, X.; Zhou, C.; Tian, Q.; Lu, Y.; et al. Genome-wide identification and
analysis of heterotic loci in three maize hybrids. Plant Biotechnol. J. 2020, 18, 185–194. [CrossRef]
Birdseye, D.; de Boer, L.A.; Bai, H.; Zhou, P.; Shen, Z.; Schmelz, E.A.; Springer, N.M.; Briggs, S.P. Plant height heterosis is
quantitatively associated with expression levels of plastid ribosomal proteins. Proc. Natl. Acad. Sci. USA 2021, 118, e2109332118.
[CrossRef]
Song, G.S.; Zhai, H.L.; Peng, Y.G.; Zhang, L.; Wei, G.; Chen, X.Y.; Xiao, Y.G.; Wang, L.; Chen, Y.J.; Wu, B.; et al. Comparative
transcriptional profiling and preliminary study on heterosis mechanism of super-hybrid rice. Mol. Plant 2010, 3, 1012–1025.
[CrossRef]
Dan, Z.; Liu, P.; Huang, W.; Zhou, W.; Yao, G.; Hu, J.; Zhu, R.; Lu, B.; Zhu, Y. Balance between a higher degree of heterosis
and increased reproductive isolation: A strategic design for breeding inter-subspecific hybrid rice. PLoS ONE 2014, 9, e93122.
[CrossRef]
Shen, G.; Hu, W.; Zhang, B.; Xing, Y. The regulatory network mediated by circadian clock genes is related to heterosis in rice. J.
Integr. Plant Biol. 2015, 57, 300–312. [CrossRef]
Zhu, D.; Zhou, G.; Xu, C.; Zhang, Q. Genetic components of heterosis for seedling traits in an elite rice hybrid analyzed using an
immortalized F2 population. J. Genet. Genom. 2016, 43, 87–97. [CrossRef]
Song, X.; Ni, Z.; Yao, Y.; Zhang, Y.; Sun, Q. Identification of differentially expressed proteins between hybrid and parents in wheat
(Triticum aestivum L.) seedling leaves. Theor. Appl. Genet. 2009, 118, 213–225. [CrossRef]
Yi, H.; Lee, J.; Song, H.; Dong, X.; Hur, Y. Genome-wide analysis of heterosis-related genes in non-heading Chinese cabbage. J.
Plant Biotechnol. 2017, 44, 208–219. [CrossRef]
Liu, T.; Duan, W.; Chen, Z.; Yuan, J.; Xiao, D.; Hou, X.; Li, Y. Enhanced photosynthetic activity in pak choi hybrids is associated
with increased grana thylakoids in chloroplasts. Plant J. 2020, 103, 2211–2224. [CrossRef] [PubMed]
Li, S.; Jayasinghege, C.P.A.; Guo, J.; Zhang, E.; Wang, X.; Xu, Z. Comparative transcriptomic analysis of gene expression
inheritance patterns associated with cabbage head heterosis. Plants 2021, 10, 275.
Li, X.; Lv, H.; Zhang, B.; Fang, Z.; Yang, L.; Zhuang, M.; Liu, Y.; Li, Z.; Wang, Y.; Zhang, Y. Dissection of two QTL clusters
underlying yield-related heterosis in the cabbage founder parent 01-20. Hortic. Plant J. 2023, 9, 77–88. [CrossRef]
Li, H.; Yuan, J.; Wu, M.; Han, Z.; Li, L.; Jiang, H.; Jia, Y.; Han, X.; Liu, M.; Sun, D.; et al. Transcriptome and DNA methylome
reveal insights into yield heterosis in the curds of broccoli (Brassica oleracea L. var. italic). BMC Plant Biol. 2018, 18, 168. [CrossRef]
Kong, X.; Chen, L.; Wei, T.; Zhou, H.; Bai, C.; Yan, X.; Miao, Z.; Xie, J.; Zhang, L. Transcriptome analysis of biological pathways
associated with heterosis in Chinese cabbage. Genomics 2020, 112, 4732–4741. [CrossRef]
Horticulturae 2023, 9, 366
17 of 18
110. Shen, Y.; Sun, S.; Hua, S.; Shen, E.; Ye, C.Y.; Cai, D.; Timko, M.P.; Zhu, Q.H.; Fan, L. Analysis of transcriptional and epigenetic
changes in hybrid vigor of allopolyploid Brassica napus uncovers key roles for small RNAs. Plant J. 2017, 91, 874–893. [CrossRef]
111. Birchler, J.A.; Yao, H.; Chudalayandi, S.; Vaiman, D.; Veitia, R.A. Heterosis. Plant Cell 2010, 22, 2105–2112. [CrossRef]
112. Wu, X.; Liu, Y.; Zhang, Y.; Gu, R. Advances in research on the mechanism of heterosis in plants. Front. Plant Sci. 2021, 12, 745726.
[CrossRef]
113. Yue, L.; Sun, R.; Li, G.; Cheng, F.; Gao, L.; Wang, Q.; Zhang, S.; Zhang, H.; Zhang, S.; Li, F. Genetic dissection of heterotic loci
associated with plant weight by Graded pool-seq in heading Chinese cabbage (Brassica rapa). Planta 2022, 255, 126. [CrossRef]
114. Choi, S.R.; Yu, X.; Dhandapani, V.; Li, X.; Wang, Z.; Lee, S.Y.; Oh, S.H.; Pang, W.; Ramchiary, N.; Hong, C.P.; et al. Integrated
analysis of leaf morphological and color traits in different populations of Chinese cabbage (Brassica rapa ssp. pekinensis). Theor.
Appl. Genet. 2017, 130, 1617–1634. [CrossRef]
115. Sun, X.; Luo, S.; Luo, L.; Wang, X.; Chen, X.; Lu, Y.; Shen, S.; Zhao, J.; Bonnema, G. Genetic analysis of Chinese cabbage reveals
correlation between rosette leaf and leafy head variation. Front. Plant Sci. 2018, 9, 1455. [CrossRef]
116. Liu, Z.; Jiang, J.; Ren, A.; Xu, X.; Zhang, H.; Zhao, T.; Jiang, X.; Sun, Y.; Li, J.; Yang, H. Heterosis and combining ability analysis of
fruit yield, early maturity, and quality in tomato. Agronomy 2021, 11, 807. [CrossRef]
117. Geleta, L.F.; Labuschagne, M.T.; Viljoen, C.D. Relationship between heterosis and genetic distance based on morphological traits
and AFLP markers in pepper. Plant Breed. 2004, 123, 467–473. [CrossRef]
118. Yang, S.; Zhang, Z.; Chen, W.; Li, X.; Zhou, S.; Liang, C.; Li, X.; Yang, B.; Zou, X.; Liu, F.; et al. Integration of mRNA and miRNA
profiling reveals the heterosis of three hybrid combinations of Capsicum annuum varieties. GM Crops Food 2021, 12, 224–241.
[CrossRef]
119. Naves, E.R.; Scossa, F.; Araújo, W.L.; Nunes-Nesi, A.; Fernie, A.R.; Zsögön, A. Heterosis for capsacinoids accumulation in chili
pepper hybrids is dependent on parent-of-origin effect. Sci. Rep. 2022, 12, 14450. [CrossRef]
120. Rodríguez-Burruezo, A.; Prohens, J.; Nuez, F. Performance of hybrids between local varieties of eggplant (Solanum melongena)
and its relation to the mean of parents and to morphological and genetic distances among parents. Eur. J. Hortic. Sci. 2008, 73, 76.
121. Liu, C.; Liu, X.; Han, Y.; Meng, H.; Cheng, Z. Heterosis prediction system based on non-additive genomic prediction models in
cucumber (Cucumis sativus L.). Sci. Hortic. 2022, 293, 110677. [CrossRef]
122. Wu, L. Relationship between SRAP marker based on genetic distance, combining ability and heterosis in pepper. Chin. J. Trop.
Crops 2020, 41, 661–668.
123. Luan, F.; Sheng, Y.; Wang, Y.; Staub, J.E. Performance of melon hybrids derived from parents of diverse geographic origins.
Euphytica 2010, 173, 1–16. [CrossRef]
124. Dafna, A.; Halperin, I.; Oren, E.; Isaacson, T.; Tzuri, G.; Meir, A.; Schaffer, A.A.; Burger, J.; Tadmor, Y.; Buckler, E.S.; et al.
Underground heterosis for yield improvement in melon. J. Exp. Bot. 2021, 72, 6205–6218. [CrossRef] [PubMed]
125. Onofri, A.; Terzaroli, N.; Russi, L. Linear models for diallel crosses: A review with R functions. Theor. Appl. Genet. 2021, 134,
585–601. [CrossRef] [PubMed]
126. Shajari, M.; Soltani, F.; Bihamta, M.R.; Alabboud, M. Genetic analysis and inheritance of floral and fruit traits in melon (Cucumis
melo) in the full diallel cross. Plant Breed. 2021, 140, 486–496. [CrossRef]
127. Aiswarya, C.S.; Vijeth, S.; Sreelathakumary, I.; Kaushik, P. Diallel analysis of chilli pepper (Capsicum annuum L.) genotypes for
morphological and fruit biochemical traits. Plants 2020, 9, 1.
128. Das, I.; Hazra, P.; Longjam, M.; Bhattacharjee, T.; Maurya, P.K.; Banerjee, S.; Chattopadhyay, A. Genetic control of reproductive
and fruit quality traits in crosses involving cultivars and induced mutants of tomato (Solanum lycopersicum L.). J. Genet. 2020, 99, 56.
[CrossRef]
129. Datta, D.R.; Rafii, M.Y.; Misran, A.; Jusoh, M.; Yusuff, O.; Haque, M.A.; Jatto, M.I. Half diallel analysis for biochemical and
morphological traits in cultivated eggplants (Solanum melongena L.). Agronomy 2021, 11, 1769. [CrossRef]
130. Kaur, S.; Sharma, S.P.; Sarao, N.K.; Deol, J.K.; Gill, R.; Abd-Elsalam, K.A.; Alghuthaymi, M.A.; Hassan, M.M.; Chawla, N.
Heterosis and combining ability for fruit yield, sweetness, β-Carotene, ascorbic acid, firmness and Fusarium wilt resistance in
muskmelon (Cucumis melo L.) involving genetic male sterile lines. Horticulturae 2022, 8, 82. [CrossRef]
131. Pavan, M.P.; Gangaprasad, S. Studies on mode of gene action for fruit quality characteristics governing shelf life in tomato
(Solanum lycopersicum L.). Sci. Hortic. 2022, 293, 110687. [CrossRef]
132. Semel, Y.; Nissenbaum, J.; Menda, N.; Zinder, M.; Krieger, U.; Issman, N.; Pleban, T.; Lippman, Z.; Gur, A.; Zamir, D. Overdominant
quantitative trait loci for yield and fitness in tomato. Proc. Natl. Acad. Sci. USA 2006, 103, 12981–12986. [CrossRef]
133. Gur, A.; Zamir, D. Unused natural variation can lift yield barriers in plant breeding. PLoS Biol. 2004, 2, e245. [CrossRef]
134. Gur, A.; Zamir, D. Mendelizing all components of a pyramid of three yield QTL in tomato. Front. Plant Sci. 2015, 6, 1096.
[CrossRef]
135. Yeager, A.F. Determinate growth in the tomato. J. Hered. 1927, 18, 263–265. [CrossRef]
136. Thouet, J.; Quinet, M.; Ormenese, S.; Kinet, J.M.; Périlleux, C. Revisiting the involvement of SELF-PRUNING in the sympodial
growth of tomato. Plant Physiol. 2008, 148, 61–64. [CrossRef]
137. Jiang, K.; Liberatore, K.L.; Park, S.J.; Alvarez, J.P.; Lippman, Z.B. Tomato yield heterosis is triggered by a dosage sensitivity of the
florigen pathway that fine-tunes shoot architecture. PLoS Genet. 2013, 9, e1004043. [CrossRef]
138. Park, S.J.; Jiang, K.; Tal, L.; Yichie, Y.; Gar, O.; Zamir, D.; Eshed, Y.; Lippman, Z.B. Optimization of crop productivity in tomato
using induced mutations in the florigen ...