Genetics of dioecy and causal sex chromosomes in plants

Journal of Genetics

Volumn 93, Issue 1, 2014, Pages 241-277

  Kumar, Sushil ^a,b   Kumari, Renu ^a,b   Sharma, Vishakha ^a  

^a SKA Institution for Research Education and Development (SKAIRED)   (India)

^b NATIONAL INSTITUTE OF PLANT GENOME RESEARCH   (India)

Author keywords

dioecy mediated species evolution; plant unisexuality; sex chromosome evolution; sex ratio variation; XY WZ transition; Y chromosome degeneration

Indexed keywords

ANIMAL; CROP; EVOLUTION; FEMALE; FLOWER; GENE AMPLIFICATION; GENETIC RECOMBINATION; GENETICS; MALE; MOLECULAR EVOLUTION; MUTATION; PLANT; PLANT CHROMOSOME; QUANTITATIVE TRAIT; REPRODUCTION; SEX CHROMOSOME; SEX DETERMINATION PROCESS; SEXUAL DEVELOPMENT;

ANIMALS; BIOLOGICAL EVOLUTION; CHROMOSOMES, PLANT; CROPS, AGRICULTURAL; EVOLUTION, MOLECULAR; FEMALE; FLOWERS; GENE AMPLIFICATION; MALE; MUTATION; PLANTS; QUANTITATIVE TRAIT, HERITABLE; RECOMBINATION, GENETIC; REPRODUCTION; SEX CHARACTERISTICS; SEX CHROMOSOMES; SEX DETERMINATION PROCESSES;

EID: 84903189544     PISSN: 00221333     EISSN: 09737731     Source Type: Journal    
DOI: 10.1007/s12041-014-0326-7     Document Type: Review

References (348)

1. Agrawal V., Sharma K., Gupta S., Kumar R., Prasad M. 2007 Identification of sex in Simmondsia chinensis (jojoba) using RAPD markers. Plant Biotechnol. Rep. 1, 207-210.
2. Ahmadi H., Bringhurst R. 1991 Genetics of sex expression in Fragaria species. Am. J. Bot. 78, 504-514.
3. Akimoto J., Fukuhara T., Kikuzawa K. 1999 Sex ratios and genetic variation in functionally androdioecious species, Sehizopepon bryoniaefolius (Cucurbitaceae). Am. J. Bot. 86, 880-886.
4. Albert A. Y. K., Otto S. P. 2005 Sexual selection can revolve sex linked sexual antagonism. Science310, 119-121.
5. Al-Dous E., George B., Al-Mahmoud M. E., Al-Jaber M. Y., Wang H., Salameh Y. M. et al. 2011 De novo genome sequencing and comparative genomics of date palm (Phoenix dactylifera). Nat. Biotech. 29, 521-527.
6. Al-Mahmoud M., Al-Dous E. K., Al-Azwani E. K., Malel J. A. 2012 DNA based essays to distinguish date palm (Arecaceae) gender. Am. J. Bot. 99, E7-E10.
7. Alexandrov O. S., Divashuk M. G., Yakovin N. A., Karlov G. I. 2012 Sex chromosome differentiation in Humulus japonicus Siebold Zuccarini, 1846 (Cannabaceae) revealed by fluorescence in situ hybridization of sub-telomeric repeat. Comp. Cytogen. 6, 239-247.
8. Allen C. E. 1930 Inheritance in a hepatic. Science71, 197-204.
9. Alstron-Rapaport C., Lascoux M., Wang Y. C., Roberts G., Tuskan G. A. 1998 Identification of RAPD marker linked to sex determination in the basket willow (Salix viminalis L.). J. Hered. 89, 44-49.
10. Ashman T. L. 2003 Constraints on the evolution of males and sexual dimorphism: field estimates of genetic architecture of reproductive traits in three population of gynodioecious Fragaria virginiana. Evolution57, 2012-2025.
11. Ashman T. L. 2005 The limits of sexual dimorphism in vegetative traits in a gynodioecious plant. Am. Nat. 166, s5-s16.
12. Atanassov I., Delichere C., Filatov D. A., Charlesworth D., Negrutiu I., and Monéger Françoise 2001 A putative monofunctional fructose-2, 6-bisphosphatase gene has functional copies located on the X and Y sex chromosomes in white campion (Silene latifolia). Mol. Biol. Evol. 18, 2162-2168.
13. Bachmann B. J. 1983 Linkage map of Escherichia coli K12, edition 7. Microbiol. Rev. 47, 180-230.
14. Bachtrog D. 2006 Expression profile of a degenerating neo-Y chromosome in Drosophila. Curr. Biol. 16, 1694-1699.
15. Bachtrog D. 2008 The temporal dynamics of processes underlying Y chromosome degeneration. Genetics179, 1513-1525.
16. Bachtrog D., Hom E., Wong K. M., Maside X., Jong P. D. 2008 Genomic degradation of a young Y chromosome in Drosophila miranda. Genome Biol. 9, R30.
17. Bachtrog D., Charlesworth B. 2002 Reduced adaptation of a non-recombining neo-Y chromosome. Nature416, 323-326.
18. Bachtrog D., Gordo I. 2004 Adaptive evolution of asexual populations under Muller's ratchet. Evol. Int. J. Org. Evol. 58, 1403-1413.
19. Bachtrog D., Kirkpatrick M., Mank J. E., McDaniel S. F., Pires J. C., Rice W., Valenzuela N. 2011 Are all sex chromosomes created equal?Trends Genet. 27, 350-357.
20. Banerjee N. S., Manoj P., Das M. R. 1999 Male-sex-associated RAPD markers in Piper longum L. Curr. Sci. 77, 693-695.
21. Barlow B. A., Weins D. 1976 Translocation heterozygosity and sex ratio in Viscum fischeri. Heredity37, 27-40.
22. Barrett S. C. H., Hough J. 2012 Sexual dimorphism in flowering plants. J. Exp. Bot. 64, 67-82.
23. Bartkowiak E. 1971 Mechanism of sex determination in Rumex hastatulus-D. Theor. Appl. Genet. 41, 320-326.
24. Bemis W. P., Wilson G. B. 1953 A new hypothesis explaining the genetics of sex determination in Spinacea oleracea L. J. Hered. 44, 91-95.
25. Bergero R., Charlesworth D. 2011 Preservation of Y-transcriptome in a 10-million-year-old plant sex chromosome system. Curr. Biol. 21, 1470-1474.
26. Bergero R., Forrest A., Kamau E., Charlesworth D. 2007 Evolutionary strata on the X-chromosome of the dioecious plant Silene latifolia, Evidence from new sex-linked genes. Genetics175, 1945-1954.
27. Bergero R., Charlesworth D., Filatov D. A., Moore R. C. 2008a Defining regions and rearrangements of the Silene latifolia Y chromosome. Genetics178, 2045-2053.
28. Bergero R., Forrest A., Charlesworth D. 2008b Active miniature transposons from a plant genome and its non-recombining Y chromosome. Genetics178, 1085-1092.
29. Bergero R., Qiu S., Forrest A., Borthwick H., Charlesworth D. 2013 Expansion of pseudoautosomal region and ongoing recombination suppression in the Silene latifolia sex chromosomes. PLoS Genet. 113, 150755.
30. Bernasconi G., Antonovics J., Biere A., Charlesworth D., Delph L. F., Filatov D. et al. 2009 Silene as a model system in ecology and evolution. Heredity103, 5-14.
31. Bhardwaj M., Eckert C. G. 2001 Functional analysis of synchronous dichogamy in flowering bush Butomus umbellatus (Butomaceae). Am. J. Bot. 88, 2204-2213.
32. Biffi R., Restivo F. M., Tassi F. 1995 A restriction fragment length polymorphism probe for early diagnosis of gender in Asparagus officinalis L. Hortic. Sci. 30, 1463-1464.
33. Birdsell J., Wills C. 1996 Significant competitive advantage confirmed by meiosis and syngamy in the yeast Saccharomyces cerevisiae. Proc. Natl. Acad. Sci. USA93, 908-918.
34. Blavet N., Blavet H., Cegan R., Zemp N., Zdanska J., Janousek B. et al. 2012 Comparative analysis of a plant pseudoautosomal region (PAR) in Silene latifolia with the corresponding S. vulgaris autosome. BMC Genomics13, 226.
35. 2 sex chromosome system and a female-biased sex ratio. Sex. Plant Reprod. 20, 171-180.
36. Bull J. J. 1983 Evolution of sex determining mechanism, pp. 316. Benjamin-Cummings, Menlo Park, USA.
37. Carroll S. B., Mulcahy D. L. 1993 Progeny sex ratios in dioecious Silene latifolia (Caryophyllaceae). Am. J. Bot. 80, 551-556.
38. Carvalho A. B. 2002 Origin and evolution of the Drosophila Y chromosome. Curr. Opin. Genet. Dev. 12, 664-668.
39. Charchar F. J., Syartman M., El-Mogharbel N., Ventura M., Kirby P., Matarazzo M. R. et al. 2003 Complex events in the evolution of the human pseudoautosomal region 2 (PAR 2). Genome Res. 13, 281-286.
40. Charlesworth D. 1984 Androdioecy and the evolution of dioecy. Biol. J. Linn. Soc. 22, 333-348.
41. Charlesworth B. 1991 The evolution of sex-chromosomes. Science251, 1030-1033.
42. Charlesworth B. 1996a The evolution of chromosomal sex determination and dasage compensation. Curr. Biol. 6, 149-162.
43. Charlesworth B. 1996b The evolution of chromosomal sex determination and dosage complementation. Curr. Biol. 6, 149-162.
44. Charlesworth B. 2002 Plant sex determination and sex chromosomes. Heredity88, 94-101.
45. Charlesworth B. 2009 Fundamental concepts in genetics: effective population size and patterns of molecular evolution and variation. Natl. Rev. Genet. 10, 195-205.
46. Charlesworth B., Charlesworth D. 1978a A model for the evolution of dioecy and gynodioecy. Am. Nat. 112, 975-997.
47. Charlesworth B., Charlesworth D. 2000 The degeneration of Y chromosomes. Philos. Trans R. Soc. Lond. B. Biol. Sci. 355, 1563-1572.
48. Charlesworth D. 2012 Plant sex chromosome evolution. J. Exp. Bot. 64, 405-420.
49. Charlesworth D. 2013 Plant sex chromosome evolution. J. Exp. Bot. 64, 405-420.
50. Charlesworth D., Charlesworth B. 1978b Population genetics of partial male-sterility and the evolution of monoecy and dioecy. Heredity41, 137-153.
51. Charlesworth D., Mank J. E. 2010 The birds and the bees and the flowers and the trees: lessons from genetic mapping of sex determination in plants and animals. Genetics186, 9-31.
52. Charlesworth D., Charlesworth B., Marais G. 2005 Steps in the evolution of heteromorphic sex chromosomes. Heredity95, 118-128.
53. Chen C. X., Yu Q., Hou S., Li Y., Eustice M., Skelton R. L. et al. 2007 Construction of a sequence-tagged high-density genetic map of papaya for comparative structural and evolutionary genomics in brassicales. Genetics177, 2481-2491.
54. Chen R. Y., Song W. Q., Li X. L. 1987 Study on the sex chromosomes of Ginkgo biloba. Proc. Sino-Jap. Symp. Plant Chromosome Res. 381-386.
55. Cherif E., Zehdi S., Castillo K., Chabrillange N., Abdoulkader S., Pintaud J. C. et al. 2013 Male specific DNA markers provide genetic evidence of an XY chromosomal system, a recombination arrest and allow the tracing of paternal lineage in date palm. New Phytol. 197, 409-415.
56. Chibalina M., Filatov D. 2011 Plant Y chromosome degeneration is retarded by haploid purifying selection. Curr. Biol. 21, 1475-1479.
57. Costantini L., Battilana J., Lamaj F., Fanizza G., Grando M. S. 2008 Berry and phenology-related traits in grapevine (Vitis Vinifera L.): From quantitative trait loci to underlying genes. BMC Plant Biol. 8, 38.
58. Costich D. E., Meagher T. R. 1992 Genetic variation in Ecballium elaterium, Breeding system and geographic distribution. J. Evol. Biol. 5: 589-601.
59. Cunado N., Navajas-Perez R, de la Herran R., Ruiz Rejon C., Ruiz Rejon M., Santos J. L., Garrido-Ramos M. A. 2007 The evolution of sex chromosomes in the genus Rumex (Polygonaceae): Identification of a new species with heteromorphic sex chromosomes. Chromosome Res. 15, 825-832.
60. Danilova T. V., Karlov G. I. 2006 Application of inter simple sequence repeat (ISSR) polymorphism for detection of sex-specific molecular markers in hop (Humulus lupulus L.). Euphytica151, 15-21.
61. De Bodt S., Maere S., van de Peer Y. 2005 Genome duplication and the origin of angiosperms. Trends Ecol. Evol. 20, 591-597.
62. Delph L. F., Arntz A. M., Scotti-Saintagne C., Scotti I. 2010 The genomic architecture of sexual dimorphism in the dioecious plant Silene latifolia. Evolution64, 2873-2886.
63. Deng C. L., Qin R. Y., Wang N. N., Cao Y., Gao J., Gao W. J., Lu L. D. 2012 Karyotype of asparagus by physical mapping of 45S and 5S rDNA by FISH. J. Genet. 91, 209-212.
64. Deputy J. C., Ming R., Ma H., Liu Z., Fitch M. M., Wang M. et al. 2002 Molecular markers for sex determination in papaya (Carica papaya L.). Theor. Appl. Genet. 106, 107-111.
65. Desfeux O., Maurice S., Henry J. P., Lejeune B., Gouyon P. H. 1996 Evolution of reproductive systems in the genus Silene. Proc. R. Soc. London, Ser. B. 263, 409-414.
66. Diggle P. K., Di Stilio V. S., Gschwend A. R., Golenberg E. M., Moore R. C., Russell J. R., Sinclair J. P. 2011 Multiple developmental processes underlie sex differentiation in angiosperm. Trends Genet. 27, 368-376.
67. Di-Stilio V. S., Kesseli R. V., Mulcahy D. L. 1998 A pseudoautosomal random amplified polymorphic DNA marker for the sex chromosomes of Silene dioica. Genetics149, 2057-2062.
68. Divashuk M. G., Alexandrov O. S., Kroupin P. Y., Karlov G. I. 2011 Molecular cytogenetic mapping of Humulus lupulus sex chromosomes. Cytogenet. Genome Res. 134, 213-219.
69. Donnison I. S., Siroky J., Vyskot B., Saedler H., Grant S. R. 1996 Isolation of Y chromosome-specific sequences from Silene latifolia and mapping of male sex-determining genes using representational difference analysis. Genetics144, 1893-1901.
70. Dorken M. E., and Barrett S. C. 2004 Sex determination and the evolution of dioecy from monoecy in Sagittaria latifolia (Alismataceae). Proc. Biol. Sci. 271, 213-219.
71. Ehsanpour A. A., Tavassoli M., Arab L. 2008 Sex determination of Pistacia vera L. using ISSR markers. Malays. Agric. Biol. 37, 25-28.
72. Elze H., Pannell J. 2011 Sexual dimorphism in androdioecious Mercurialis annua, a wind-pollinated herb. Int. J. Plant Sci. 172, 49-59.
73. Eppley S. M., Pannell J. R. 2007 Density-dependent self-fertilization and male versus hermaphrodite siring success in an androdioecious plant. Evolution61, 2349-2359.
74. Eppley S. M., Maureen L. S., Grosberg R. K. 1998 Intrapopulation sex ratio variation in the salt grass Distichlis spicata. Am. Nat. 152, 659-670.
75. Ezaz T., Quinn A. E., Miura I., Sarre S. D., Georges A., Marshall Graves J. A. 2005 The dragon lizard Pogna utticeps has ZZ / ZW micro-sex chromosomes. Chromosome Res. 13, 763-766.
76. Ezaz T., Stiglec R., Veyrunes F., Graves J. A. M. 2006 Relationships between vertebrate ZW and XY sex chromosome systems. Curr. Biol. 16, R736-R743.
77. Ezaz T., Quinn A. E., Sarre S. D., O'Meally D., Georges A., Marshall G. J. A. 2009a Molecular marker suggests rapid changes in sex-determining mechanisms in Australian dragon lizards. Chromosome Res17, 91-98.
78. Ezaz T., Moritz B., Waters P., Marshall Graves J. A., Georges A., Sarre S. D. 2009b The ZW sex microchromosomes of an Australian dragon lizard share no homology with those of other reptiles or birds. Chrom. Res. 17, 965-973.
79. Farbos I., Veuskens J., Vyskot B., Oliveira M., Hinnisdaels S., Aghmir, A. et al. 1999 Sexual dimorphism in white campion: deletion on the Y chromosome results in a floral asexual phenotype. Genetics151, 1187-1196.
80. Felsenstein J. 1974 The evolutionary advantage of recombination. Genetics78, 737-756.
81. Ferris P. J., Armbrust E. V., Goodenough V. 2002 Genetic structure of the mating-type locus of Chlamydomonas reinhardtii. Genetics160, 181-200.
82. Field D. L., Pickup M., Barrett S. C. 2012 Comparative analyses of sex-ratio variation in dioecious plants. Evolution67, 660-672.
83. Field D. L., Pickup M., Barrett S. C. 2013 Ecological context and metapopulation dynamics affect sex-ratio variation among dioecious plant populations. Ann. Bot. 111, 917-923.
84. Filatov D. A. 2005a Evolutionary history of Silene latifolia sex chromosomes revealed by genetic mapping of four genes. Genetics170, 975-979.
85. Filatov D. A. 2005b Substitution rates in a new Silene latifolia sex-linked gene, SlssX/Y. Mol. Biol. Evol. 22, 402-408.
86. Filatov D. A., Charlesworth D. 2002 Substitution rates in the X- and Y- linked genes of the plants, Silene latifolia and S. dioica. Mol. Biol. Evol. 19, 898-907.
87. Filatov D. A., Moneger F., Negrutiu I., Charlesworth D. 2000 Evolution of a plant Y-chromosome: Variability in a Y-linked gene of Silene latifolia. Nature404, 388-390.
88. Filatov D. A., Laporte V., Vitte C., Charlesworth D. 2001 DNA diversity in sex-linked and autosomal genes of the plant species Silene latifolia and Silene dioica. Mol. Biol. Evol. 18, 1442-1454.
89. Flores-Renteria L., Molina-Freaner F., Whipple A. V., Gehring C. A., Dominguez C. A. 2013 Sexual stability in the nearly dioecious Pinus johannis (Pinaceae). Am. J. Bot. 100, 602-612.
90. Frajman B., Eggens F., Oxelman B. 2009 Hybrid origins and homoploid reticulate evolution within Heliosperma (Sileneae, Caryophyllaceae)- a multigens phylogenetic approach with relative dating. Syst. Biol. Biol. 58, 328-345.
91. Franchini P., Colangelo P., Solano E., Capanna E., Verheyen E. and Castiglia R. 2010 Reduced gene flow at pericentromeric loci in a hybrid zone involving chromosomal races of the house mouse Mus musculus domesticus. Evolution64, 2020-2032.
92. Fraser J. A., Heitman J. 2004 Evolution of fungal sex chromosomes. Mol. Microbiol. 51, 299-306.
93. Fraser L. G., Tsang G. K., Datson P. M., De Silva H. N., Harvey C. F., Gill G. P. et al. 2009 A gene-rich linkage map in the dioecious species Actinidia chinensis (kiwifruit) reveals putative X/Y sex determining chromosomes. BMC Genomics10, 102.
94. Freeman D. C., Wachocki B. A., Stender M. J., Goldschlag D. E., Michaels H. J. 1994 Seed size and sex ratio in spinach, application of the Trivers-Willard hypothesis to plants. Ecoscience1, 54-63.
95. Fujisawa M., Hayashi K., Nishio T., Bando T., Okada S., Yamato K. T., Fukuzawa H. and Ohyama K. 2001 Isolation of X and Y chromosome-specific DNA markers from a liverwort, Marchantia polymorpha, by representational difference analysis. Genetics159, 981-985.
96. Galan F. 1946 Sur la genetique de la monoecie et la dioecie zygotique chez Ecballium eletarium Rich. Crit. Rev. Hebd. Seances Acad. Sci. Paris222, 1130-1131.
97. Gangopadhyay G., Roy S. K., Ghose K., Poddar R., Bandyopadhyay, T., Basu D., Mukherjee K. K. 2007 Sex detection of Carica papaya and Cycas circinalis in pre-flowering stage by ISSR and RAPD. Curr. Sci. 92, 524-526.
98. Gao W. J., Li R. L., Li S. F., Deng C. L., Li S. P. 2007 Identification of two markers linked to the sex locus in dioecious Asparagus officinalis plants. Russ. J. Plant Physiol. 54, 816-821.
99. Gaut B. S. 1998 Molecular clocks and nucleotide substitution rates in higher plants. Evol. Biol. 30, 93-100.
100. George J., Karun A., Manimekalai R., Rajesh M. K., Remya P. 2007 Identification of RAPD markers linked to sex determination in palmyrah (Borassus flabellifer L.). Curr. Sci. 93, 1075-1077.
101. Gill G. P., Harvey C. F., Gardner R. C., Fraser L. G. 1998 Development of sex-linked PCR markers for gender identification in Actinidia. Theor. Appl. Genet. 97, 439-445.
102. Goddard M. R., Godfray H. C. J., Burt A. 2005 Sex increases the efficacy of natural selection in experimental yeast populations. Nature434, 636-640.
103. Goldberg M. T., Spigler R. B., Ashman T.-L. 2010 Comparative genetic mapping points to different sex chromosomes in sibling species of wild strawberry (Fragaria). Genetics156, 1425-1433.
104. Gorelick R. 2005 Theory for why dioecious plants have equal length chromosomes. Am. J. Bot. 92, 979-984.
105. Grabowska-Joachimiak A., Jochimiak A. 2002 C-banded karyotypes of two Silene species with heteromorphic sex chromosomes. Genome45, 243-252.
106. Grabowska-Joachimiak A., Seliwinska E., Pigula M., Skomra U., Joachimiak A. J. 2006 Genome size in Humulus lupulus L. and H. japonicus Siebold and Zucc (Cannabaceae). Acta. Soc. Bot. Pol. 75, 207-214.
107. Grabowska-Joachimiak A., Mosiolek M., Lech A., Goralski G. 2011 C-banding/DAP1 and in situ hybridization reflect karyotype structure and sex chromosome differentiation in Humulus japonicus Siebold and Zucc. Cytogenet. Genome Res. 132, 203-211.
108. Grant S., Hunkirchen B., Saedler H. 1994a Developmental differences between male and female flowers in the dioecious plant Silene latifolia. Plant J. 6, 471-480.
109. Grant S., Siroky J., Pan W.-H., Macas J., Saedler H. 1994b Genetics of sex determination in flowering plants. Develop. Genet. 15, 214-230.
110. Graves J. A. M. 2006 Sex chromosome specialization and degeneration in mammals. Cell24, 901-914.
111. Graves J. A. M., Peichel C. L. 2010 Are homologies in vertebrate sex determination due to shared ancestory or to limited options? Genome Biol. 11, 205.
112. Greilhuber J., Borsch T., Muller K., Worberg A., Porembski S., Barthlott W. 2006 Smallest angiosperm genomes found in Lentibulariaceae with chromosomes of bacterial size. Plant Biol. 8, 770-777.
113. Greilhauber J., Leitch I. J. 2013 Plant genome diversity. In Physical structure, behaviour and evolution of plant genomes (ed. I. J. Leitch, J. Greilhuber, J. Dolezel and J. F. Wendel), vol 2, pp. 323-344. Springer-Verlag, Wien.
114. Gschwend A., Yu Q., Tong E. J., Zeng F., Han J., VanBuren R. et al. 2012 Rapid divergence and expansion of the X chromosome in papaya. Proc. Natl. Acad. Sci. USA109, 13716-13721.
115. Gunter L. E., Roberts G. T., Lee K., Larimer F. W., Tuskan G. A. 2003 The development of two flanking SCAR markers linked to a sex determination locus in Salix viminalis. J. Hered. 94, 185-189.
116. Hair J. B., Beuzenberg E. J. 1958 Chromosome evolution in Podocarpaceae. Nature181, 1584-1586.
117. Hamilton W. D. 1967 Extraordinary sex ratios. Science156, 477-488.
118. Harris M. S., Pannell J. R. 2010 Canopy seed storage is associated with sexual dimorphism in the woody dioecious genes Leucadendron. J. Ecol. 98, 509-515.
119. Harvey C. F., Gill G. P., Fraser L. G., McNeilage M. A. 1997 Sex determination in Actinidia. 1. Sex-linked markers and progeny sex ratio in diploid A. chinensis. Sex Plant Reprod. 10, 149-154.
120. Heilbuth J. C. 2000 Lower species richness in dioecious clades. Am. Nat. 156, 221-241.
121. Heilbuth J. C., Ilves K. L., Otto S. P. 2001 The consequences of dioecy for seed dispersal: modeling the seed-shadow handicap. Evolution55, 880-888.
122. Hillis D. M., Green D. M. 1990 Evolutionary changes of heterogametic sex in the phylogenetic history of amphibians. J. Evol. Biol. 3, 49-64.
123. Hizume M., Shiraishi H., Tanaka A. 1988 A cytological study of Podocarpus macrophyllus with special reference to sex chromosomes. Jap. J. Genet. 63, 413-423.
124. Hobza R., Lengerova M., Svoboda J., Kubekova H., Kejnovsky E., Vyskot B. 2006 An accumulation of tandem DNA repeats on the Y chromosome in Silene latifolia during early stages of sex chromosome evolution. Chromosoma115, 376-382.
125. Hobza R., Kejnovsky E., Vyskot B., Widmer A. 2007 The role of chromosomal rearrangements in the evolution of Silene latifolia sex chromosomes. Mol. Genet. Genomics278, 633-638.
126. Hofmeyr J. D. J. 1938 Genetic studies of Carica papaya L. S. Afr. J. Sci. 35, 300-304.
127. Holstein N. 2012 Evolution, biogeography, and monographic treatment of Coccinia (Cucurbitaceae). Ph. D. thesis, Ludwig-Maxmilians University, Munich, Germany.
128. Holstein N., Renner S. S. 2011 Niche conservation? Biome switching within and between species of the African genus Coccinia (Cucurbitaceae). BMC Evol. Biol. 11, 28.
129. Honys D., Twell D. 2004 Transcriptome analysis of haploid male gametophyte development in Arabidopsis. Genome Biol. 5, 85.
130. Hormuza J. I., Dollo L., Polito V. S. 1994 Identification of RAPD marker linked to sex determination in Pistacia vera using bulked segregant analysis. Theor. Appl. Genet. 89, 9-13.
131. Hou W., Fan J., Zhou F., Zhao S. 2009 RAPD markers related to sex locus in Populus tomentose. Front. For. Chin4, 223-226.
132. Hough J., Immler S., Barrett C. H., Otto S. P. 2013 Evolutionarily stable sex ratios and mutation load. Evolution67, 1915-1925.
133. Howell E. C., Armstrong S. J., Filatov D. A. 2009a Evolution of neo-sex chromosomes in Silene diclinis. Genetics182, 1109-1115.
134. Howell E. C., Armstrong S., Filatov D. 2009b Evolution of neo-sex chromosomes in Silene diclinis. Genetics182, 1109-1115.
135. Hussaini F. S., Hassani H. S., Arvin M. J., Baghizadeh A., Mohammadi-Nejad G. 2011 Sex determination of jojoba (Simmondsia chinensis cv Arizona) by random amplified polymorphic DNA (RAPD) molecular markers. Afr. J. Biotech. 10, 470-474.
136. Ironside J. E., Filatov D. A. 2005 Extreme population structure and high interspecific divergence of the Silene Y-chromosome. Genetics171, 705-713.
137. Ishizaki K., Ishizaki K., Shimizu-Ueda Y., Okada S., Yamamoto M., Fujisawa M. et al. 2002 Multicopy genes uniquely amplified in the Y-chromosome-specific repeats of the liverwort Marchantia polymorpha. Nucleic Acids Res. 30, 4675-4681.
138. Jaarola M., Martin R. H., Ashley T. 1998 Direct evidence for suppression of recombination within two pericentric inversions in humans: a new sperm-fish technique. Am. J. Hum. Genet. 63, 218-224.
139. Jakse J., Stajner N., Kozjak P., Cerenak A., Javornik B. 2008 Trinucleotide micro-satellite repeat is tightly linked to male sex in hop (Humulus lupulus L.)Mol. Breed. 21, 139-148.
140. Jamilena M., Marriotti B., Manzano S. 2008 Plant sex chromosomes: molecular structure and function. Cytogenet. Genome Res. 120, 255-264.
141. Jiang C., Sink K. C. 1997 RAPD and SCAR markers linked to the sex expression locus M in asparagus. Euphytica94, 329-333.
142. Kafer J., Talianova M., Bigot T., Michu E., Gueguen L., Widmer A. et al. 2013 Patterns of molecular evolution in dioecious and non-dioecious Silene. J. Evol. Biol. 26, 335-346.
143. Kaiser V. B., Bachtrog D. 2010 Evolution of sex chromosomes in insects. Annu. Rev. Genet. 44, 91-112.
144. Kaiser V. B., Bergero R., Charlesworth D. 2009 Slcyt, a newly identified sex-linked gene, has recently moved onto the X chromosome in Silene latifolia (Caryophyllaceae). Mol. Biol. Evol. 26, 2343-2351.
145. Kaltz O., Bell G. 2002 The ecology and genetics of fitness in Chlamydomonas XII. Repeated sexual episodes increase rate of adaptation to novel environments. Evolution56, 1743-1753.
146. Karlov G. L., Danilova T. V., Horlemann C., Weber G. 2003 Molecular cytogenetic in hop (Humulus lupulus L.) and identification of sex chromosomes by DAPI-banding. Euphytica132, 185-190.
147. Kater M. M., Franken J., Carney K. J., Colombo L., Angenent G. C. 2001 Sex determination in the monoecious cucumber is confined to specific floral whorls. Plant Cell13, 481-493.
148. Kejnovsky E., Vrana J., Matsunaga S., Soucek P., Siroky J., Dolezel J., Vyskot B. 2001 Localization of male-specifically expressed MROS genes of Silene latifolia by PCR on flow-sorted sex chromosomes and autosomes. Genetics158, 1269-1277.
149. Khadka D. K., Nejidat A., Tal M., Golan-Goldhirsh A. 2002 DNA markers for sex: molecular evidence for gender dimorphism in dioecious Mercurialis annua L. Mol. Breed. 9, 251-257.
150. Khadka D. K., Nejidat A., Tal M., Golan-Goldhirsh A. 2005 Molecular characterization of a gender-linked DNA marker and a related gene in Mercurialis annua L. Planta222, 1063-1070.
151. Khattak J. Z. K., Torp A. M., Andersen S. B. 2006 A genetic linkage map of Spinacea oleracea and localization of a sex determination locus. Euphytica148, 311-318.
152. Kihara H., Ono T. 1923 Cytological studies of Rumex L. Bot. Mag. 37, 84-90.
153. Kim S.-Y., Kim C.-S., Lee J., Bang J.-W. 2008 Karyotype analysis and physical mapping using two rRNA genes in dioecious plant, Humulus japonicus Sieboid and Zucc. Genes Genomics30, 157-161.
154. Korpelainen H., Bisang I., Hedenas L., Kolehmainen J. 2008 The first sex-specific molecular marker discovered in the moss Pseudocalliergon trifarium. J. Hered. 99, 581-587.
155. Krizek B. A., Fletcher J. C. 2005 Molecular mechanisms of flower development: An Armchair guide. Nat. Rev. Genet6, 688-698.
156. Kumar L. S. S., Viseveshwaraiah S. 1952 Sex mechanism in Coccinia indica Wight and Arn. Nature170, 330-331.
157. Kumar S. 2007 Conventional and new concepts of species and genetic mechanisms for the origin of species. Proc. Ind. Natl. Sci. Acad. 73, 111-120.
158. Kumar S., Singh B. D., Sinha D. P., Rai M. 2008 Sex expression-associated RAPD markers in pointed gourd (Trichosanthes dioica). In Proceedings of the IXth EUCARPIA meeting on genetics and breeding of Cucurbitaceae (ed. M. Pitrat), pp. 543-550. Avignon, France.
159. Kumar S., Kumari R., Sharma V., Sharma V. 2013a Roles and establishment, maintenance and erasing of the epigenetic cytosine methylation marks in plants. J. Genet. 92, 629-666.
160. Kumar S., Kumari R., Sharma V., Yadav G. 2013b Common and distinguishing characteristics of genes and genomes and their evolution in the genome sequenced legumes. Proc. Ind. Natl. Sci. Acad. 79, 277-286.
161. Kurita M., Kuroki Y. 1970 Y-chromosome and heterochromatin in Rumex acetosa. Jpn. J. Genet. 45, 255-260.
162. Lan T., Zhang S., Liu B., Li X., Chen R., Song W. 2006 Differentiating sex chromosomes of the dioecious Spinacia oleracea L. (spinach) by FISH of 45S rDNA. Cytogenet. Genome Res. 114, 175-177.
163. Lan T., Chen R. Y., Li X. L., Dong F. P., Qi Y. C., Song W. C. 2008 Microdissection and painting of the W chromosome in Ginkago biloba showed different labelling patterns. Bot. Stud. 49, 33-37.
164. Lardon A., Georgiev S., Aghmir A., Le Merrer G., Negrutiu I. 1999 Sexual dimorphism in white campion: complex control of carpel number is revealed by Y chromosome deletions. Genetics151, 1173-1185.
165. Laporte V., Filatov D. A., Kamau E., Charlesworth D. 2005 Indirect evidence from DNA sequence diversity for genetic degeneration of the Y-chromosome in dioecious species of the plant Silene, the Sly4/Slx4 and DD44-X/DD44-Y gene pairs. J. Evol. Biol. 18: 337-347.
166. Lebel-Hardenack S., Hauser E., Law T. F., Schmid J., Grant S. R. 2002 Mapping of sex determination loci on the white campion (Silene latifolia) Y chromosome using amplified fragment length polymorphism. Genetics160, 717-725.
167. Lederberg J., Tatum E. L. 1946 Gene recombination in E. coli. Nature158, 558.
168. Lengerova M., Moore R. C., Grant S. R., Vyskot B. 2003 The sex chromosomes of Silene latifolia revisited and revised. Genetics165, 935-938.
169. Lengerova M., Kejnovsky E., Hobza R., Macas J., Grant S. R., Vyskot B. 2004 Multicolor FISH mapping of the dioecious model plant, Silene latifolia. Theor. Appl. Genet. 108, 1193-1199.
170. Lewis D 1942 The evolution of sex in flowering plants. Biolog. Rev. 17, 46-67.
171. Liao L., Liu J., Dai Y., Li Q., Xie M., Chen Q. et al. 2009 Development and application of SCAR markers for sex identification in the dioecious species Ginkgo biloba L. Euphytica169, 49-55.
172. Lindholm A., Breden F. 2002 Sex chromosomes and sexual selection in poeciliid fishes. Am. Nat. 160, 214-224.
173. Liu Z., Moore P. H., Ma H., Ackerman C. M., Ragiba M., Yu Q., Pearl H. M. 2004 A primititive Y chromosome in papaya marks incipient sex chromosome evolution. Nature427, 348-352.
174. Loptien H. 1979 Identification of the sex chromosome pair in asparagus (Asparagus officinalis L.). Z. Pflanzenzuecht. 82, 162-173.
175. Ma H., Moore P. H., Liu Z., Kim M. S., Yu Q., Fitch M. M. M. et al. 2004 High-density linkage mapping revealed suppression of recombination at the sex determination locus in papaya. Genetics166, 419-436.
176. Macas J., Kejnovsky E., Neumann P., Novak P., Koblizkova A., Vyskot B. 2008 Next generation sequencing-based analysis of repetitive DNA in the model dioecious plant Silene latifolia. PLoS One6, e27335.
177. Magdum S., Banerjee U., Murugan P., Gangapur D., Ravikesvan R. 2013 Gene duplication as a major force in evolution. J. Genet. 92, 155-161.
178. Mandolino G., Carboni A., Forapani S., Faeti V., Ranalli P. 1999 Identification of DNA markers linked to the male sex in dioecious hemp (Cannabis sativa L.). Theor. Appl. Genet. 98, 86-92.
179. Mank J. E., Promislo D. E. L., Avise J. C. 2006 Evolution of alternative sex-determining mechanism in teleost fishes. Biol. J. Linn. Soc. 87, 83-93.
180. Mank J. E., Vicoso B., Berlin S., Charlesworth B. 2010 Effective population size and the faster-X effect: empirical results and their interpretation. Evolution64, 663-674.
181. Manoj P., Banerjee N. S., Ravichandran P. 2005 Development of sex-associated SCAR markers in Piper longum L. PGR Newslett. 141, 44-50.
182. Marais G. A. B., Nicolas M., Bergero R., Chambrier P., Kejnovsky E., Moneger F. et. al. 2008 Evidence for degeneration of the Y-chromosome in the dioecious plant Silene latifolia. Curr. Biol. 18, 545-549.
183. Marais G. A. B., Forrest A., Kamau E., Kafer J., Daubin V., Charlesworth D. 2011 Multiple nuclear gene phylogenetic analysis of the evolution of dioecy and sex chromosomes in the genus Silene. PLoS One6, e21915.
184. Marguerit E., Boury C., Manicki A., Donnart M., Butterlin G., Nemorin A., Wiedemann-Merdinoglu S. et al. 2009 Genetic dissection of sex determinism, inflorescence morphology and downy mildew resistance in grapevine. Theor. Appl. Genet. 118, 1261-1278.
185. Marin I., Siegal M. L., Baker B. S. 2000 The evolution of dosage-compensation mechanisms. BioEssays22, 1106-1114.
186. Mariotti B., Manzano S., Kejnovsky E., Vyskot B., Jamilena M. 2009 Accumulation of Y-specific satellite DNAs during the evolution of Rumex acetosa sex chromosome. Mol. Genet. Genomics281, 249-259.
187. Mariziani G., Caporali E. and Spada A. 1999 Search for genes involved in asparagus sex-determination. In Sex determination in plants (ed. C. C. Ainsworth) pp. 149-162. Bios Scientific Publishers, Oxford, UK.
188. Martin A., Troadec C., Boualem A., Rajab M., Fernandez R., Morin H. et al. 2009 A transposon-induced epigenetic change leads to sex determination in melon. Nature461, 1135-1138.
189. Martin F. W. 1966 Sex ratio and sex determination in Dioscorea. J. Hered57, 95-99.
190. Mather K. 1950 The genetical architecture of heterostyly in Primula sinensis. Evolution4, 340-352.
191. Matsubara K., Tarui H., Toriba M., Yamada K., Nishida-Umehara C., Agata K., Matsuda Y. 2006 Evidence for different origin of sex chromosomes in snakes, birds and mammals and step-wise differentiation of snake sex chromosomes. Proc. Natl. Acad. Sci. USA103, 18190-19195.
192. McDaniel S. F. 2005 Genetic correlations do not constrain the evolution of sexual dimorphism in the moss Ceratodon purpureus. Evolution59, 2353-2361.
193. McDaniel S. F., Atwood J., Burleigh J. G. 2012 Recurrent evolution of dioecy in bryophytes. Evolution67, 567-572.
194. McDaniel S. F., Willis J. H., Shaw A. J. 2007 A linkage map reveals a complex basis for segregation distortion in an interpopulation cross in the moss Ceratodon purpureus. Genetics176, 2489-2500.
195. McLetchie D. N., Collius A. L. 2001 Identification of DNA regions specific to the X and Y chromosomes of Sphaerocarpus taxanus. Bryologist104, 543-547.
196. Mendez M., Munzinger J. 2010 Planchonella, first record of gynomonoecy for the family Sapotaceae. Plant. Syst. Evol. 287, 65-73.
197. Midgley J. J. 2010 Causes of secondary sexual differences in plants-evidence from extreme leaf dimorphism in Leucadendron (Priteaceae). South Afr. J. Bot76, 588-592.
198. Milewicz M., Sawicki J. 2013 Sex linked markers in dioecious plants. POJ6, 144-149.
199. Miller J. S., Venable D. L. 2000 Polyploidy and the evolution of gender dimorphism in plants. Science289, 2335-2338.
200. Ming R., Hou S., Feng Y., Yu Q., Dionne-Laporte A., Saw J. H. et al. 2008 The draft genome of the transgenic tropical fruit tree papaya (Carica papaya Linnaeus). Nature452, 991-996.
201. Ming R., Bendahmane A., Renner S. S. 2011 Sex chromosomes in land plants. Ann. Rev. Plant Biol. 62, 485-574.
202. Mitchell C. H., Diggle P. K. 2005 Evolution of unisexual flowers: morphological and functional convergence results from diverse developmental transitions. Am. J. Bot. 92, 1068-1076.
203. Moore R. C., Kozyreva O., Lebel-Hardenack S., Siroky J., Hobza R., Vyskot B., Gran S. R. 2003 Genetic and functional analysis of DD44, a sex-linked gene from the dioecious plant Silene latifolia, provides clues to early events in sex chromosome evolution. Genetics163, 321-334.
204. Mrackova M., Nicolas M., Hobza R., Negrutiu I., Moneger F., Widmer, A. et al. 2008 Independent origin of sex chromosomes in two species of genus Silene. Genetics179, 1129-1133.
205. Muller H. J. 1964 The relation of recombination to mutational advance. Mut. Res. 106, 2-9.
206. Murray M. J. 1940 The genetics of sex determination in the family Amaranthaceae. Genetics35, 409-431.
207. Muyle A., Zemp N., Deschamps C., Mousset S., Widmer A. et al. 2012 Rapid de novo evolution of X chromosome dosage compensation in Silene latifolia, a plant with young sex chromosomes. PLoS Biol. 10, e1001308.
208. Mwase W. F., Erik-Lid S., BjOrnstad A., Stedje B., Kwapata M. B., Bokosi J. M. 2007 Application of amplified fragment length polymorphism (AFLPs) for detection of sex-specific markers in dioecious Uapaca kirkiana Muell Arg. Afr. J. Biotechnol. 6, 137-142.
209. Na J. K., Wang J., Murray J. E., Gschwend A. R., Zhang W., Yu Q. et al. 2012 Construction of physical maps for the sex-specific regions of papaya sex chromosomes. BMC Genomics13, 176.
210. Naurin S., Hansson B., Bensch S., Hesselquist D. 2009 Why does dosage compensation differ between XY and ZW taxa?Trends Genet26, 15-20.
211. Navajas-Perez R., la Herran Rd, Jamilena M., Lozano R., Rejon C. R, Rejon M. R., Garrido-Ramos M. A. 2005a Reduced rates of sequence evolution of Y-linked satellite DNA in Rumex (Polygonaceae). J. Mol. Evol. 60, 391-399.
212. Navajas-Perez R., de la Herran R., Lopez Gonzalez G., Jamilena M., Lozano R., Ruiz Rejon C. et al. 2005b The evolution of reproductive systems and sex determining mechanisms within Rumex (Polygonaceae) inferred from nuclear and chloroplastidial sequence data. Mol. Biol. Evol. 22, 1929-1939.
213. Navajas-Perez R., Schwarzacher T, de la Herran R., Ruiz Rejon C., Ruiz Rejon M., Garrido-Ramos M. A 2006 The origin and evolution of the variability in a Y-specific satellite-DNA of Rumex acetosa and its relatives. Gene368, 61-71.
214. Navajas-Perez R., Schwarzacher T., Rejon M. R., Garrido-Ramos M. A. 2009 Molecular cytogenetic characterization of Rumex papillaris, a dioecious plant with an XX/XY(1)Y2 sex chromosome system. Genetica135, 87-93.
215. Negrutiu I., Vyskot B., Barbacar N., Georgiev S., Moneger F. 2001 Dioecious plants. A key to the early events of sex chromosome evolution. Plant Physiol. 127, 1418-1424.
216. Nicolas M., Marais G., Hykelova V., Janousek B., Laporte V., Vyskot B. et al. 2005 A gradual process of recombination restriction in the evolutionary history of the sex chromosomes in dioecious plants. PLoS Biol. 3, 47-56.
217. Nilsson E., Agren J. 2006 Population size, female fecundity, and sex ratio variation in gynodioecious Plantago maritima. J. Evol. Biol. 19, 825-833.
218. Nishiyama R., Ishii K., Kifune E., Kazama Y., Nishihara K., Matsunaga S. et al. 2010 Sex chromosome evolution revealed by physical mapping of SIAP3X/Y in the dioecious plants Silene latifolia. Cytologia75, 319-325.
219. Obbard D. J., Harris S. A., Buggs R. J., Pannell J. R. 2006 Hybridization, polyploidy and the evolution of sexual systems in Mercurialis (Euphorbiaceae). Evolution60, 1801-1815.
220. Ohno S. 1967 Sex chromosomes and sex-linked genes. In Monographs on endocrinology, vol. 1, pp. 192. Springer, Berlin.
221. Okada S., Fujisawa M., Sone T., Nakayama S., Nishiyama R., Takenaka, M. et al. 2000 Construction of male and female PAC genomic libraries suitable for identification of Y-chromosome-specific clones from liverwort, Marchantia polymorpha. Plant J. 24, 421-428.
222. Okada S., Sone T., Fujisawa M., Nakayama S., Takenaka M., Ishizaki K. et al. 2001 The Y chromosome in the liverwort Marchantia polymorpha has accumulated unique repeat sequences harbouring a male-specific gene. Proc. Natl. Acad. Sci. USA98, 9454-9459.
223. Ono T. 1955 Studies on hop. I. Chromosomes of common hop and its relatives. Bull. Brew. Sci. 2, 1-65.
224. Onodera Y., Yonaha I., Masumo H., Tanaka A., Niikura S., Yamazaki S., Mikami T. 2011 Mapping of the genes for dioecism and monoecism in Spinacia oleracea L.: evidence that both genes are closely linked. Plant Cell Rep. 30, 965-971.
225. Otto S. P., Lenormand T. 2002 Resolving the paradox of sex and recombination. Natl. Rev. Genet. 3, 252-261.
226. Otto S. P., Pannell J. R., Peichel C. L., Ashman T. L., Charlesworth D., Chippindale A. K. et al. 2011 About PAR: The distinct evolutionary dynamics of the pseudoautosomal region. Trends Genet. 27, 358-367.
227. Oyama R. K., Volz S. M., Renner S. S. 2009 A sex-linked SCAR marker in Bryonia dioica (Cucurbitaceae); a dioecious species with XY sex-determination and homomorphic sex chromosomes. J. Evol. Biol. 22, 214-224.
228. Oyama R. K., Silber M. V., Renner S. S. 2010 A specific insertion of a solo-LTR characterizes the Y-chromosome of Bryonia dioica (Cucurbitaceae). BMC Res. Notes3, 166-173.
229. Pakull B., Groppe K., Meyer M., Markussen T., Fladung M. 2009 Genetic linkage mapping in aspen (Populus tremula L. and Populus tremuloidex Michx). Tree Genet. Genomics5, 505-515.
230. Pannell J. 1997a Mixed genetic and environmental sex determination in an androdioecious population of Mercurialis annua. Heredity78, 50-56.
231. Pannell J. R. 1997b Widespread functional androdioecy in Mercurialis annua L. (Euphorbiaceae). Biol. J. Linn. Soc. 61, 95-116.
232. Pannell J. R. 2008 Gender variation and transitions between sexual systems in Mercurialis annua. Int. J. Plant Sci. 169, 129-139.
233. Pannell J. R., Obbard D. J., Buggs R. J. A. 2004 Polyploidy and the sexual system: What can we learn from Mercurialis annua. Biol. J. Linn. Soc. 82, 547-560.
234. Parasnis A. S., Ramakrishna W., Chowdari K. V., Gupta V. S., Ranjekar P. K. 1999 Microsatellite (GATA)n reveals sex specific differences in papaya. Theor. Appl. Genet. 99, 1047-1052.
235. Parasnis A. S., Gupta V. S., Tamhankar S. A., Ranjekar P. K. 2000 A highly reliable sex diagnostic PCR assays for mass screening of papaya seedlings. Mol. Breed. 6, 337-344.
236. Patel G. I. 1952 Chromosome basis of dioecism in Trichosanthes dioica Roxb. Curr. Sci. 21, 343-344.
237. Paolucci I., Gaudet M., Jorge V., Isacco Beritognolo I., Terzoli S., Kuzminsky E. et al. 2008 Genetic linkage maps of Populus alba L. and comparative mapping analysis of sex determination across Populus species. Tree Genet. Genomics6, 863-875.
238. Peiffer J. A., Kaushik S., Sakai H., Arteaga-Vazquez M., Sanchez-Leon N., Ghazal H. et al. 2008 A spatial dissection of the Arabidopsis floral transcriptome by MPSS. BMC Plant Biol. 8, 43.
239. Peil A., Flachowsky H., Schumann E., Weber W. E. 2003 Sex-linked AFLP markers indicate a pseudoautosomal region in hemp (Cannabis sativa L.)Theor. Appl. Genet. 107, 102-109.
240. Persson H. A., Nybom H. 1998 Genetic sex determination and RAPD marker segregation in the dioecious species sea buckthorn (Hippophoe rhamnoides L.). Hereditas129, 45-51.
241. Polley A., Ganal M. W., Seigner E. 1997 Identification of sex in hop (Humulus lupulus) using molecular markers. Genome40, 357-361.
242. Prakash S., Staden J. V. 2006 Sex identification in Encephalartos natalensis (Dyer and Verdoorn) using RAPD markers. Euphytica152, 197-200.
243. Price T. D. 1984 The evolution of sexual size dimorphism in Darwin's finches. Am. Nat. 123, 500-518.
244. Prince E. G., Kirkland D., Demutti J. P. 2010 Hyperexpression of the X chromosome in both sexes results in extensive female bias of X-linked genes in the flour beetle. Genome Biol. Evol. 2, 336-346.
245. Pritham E. J., Zhang Y. H., Feschotte C., Kesseli R. V. 2003 An Ac-like transposable element family with transcriptionally active Y-linked copies in the white campion Silene latifolia. Genetics165, 799-807.
246. Qiu Y. L., Palmer J. D. 1999 Phylogeny of early land plants: insights from genes and genomes. Trends Plant Sci. 4, 26-30.
247. Quinn A. E., Georges A., Sarre S. D., Guarino F., Ezaz T., Graves J. A. 2007 Temperature sex reversal implies sex gene dosage in a reptile. Science316, 411.
248. Rahman M. A., Ainsworth C. C. 2004 AFLP analysis of genome difference between males and females in dioecious plant Rumex acetosa. J. Bio. Sci. 4, 160-169.
249. Rai L. K., Singh K. K. 2013 Phoenix rupicola in the Eastern Himalaya. Curr. Sci. 104, 572-573.
250. Ranker T. A., Haufler C. H. 2008 Biology and evolution of ferns and lycophytes. Cambridge University Press, Cambridge, UK.
251. Rautenberg A., Hathaway L., Oxelman B., Prentice H. C. 2010 Geographical phylogenetic patterns in Silene section Melandrium (Caryophyllaceae) as inferred from chloroplast and nuclear DNA sequences. Mol. Phylogenet. Evol. 57, 978-991.
252. Rautenberg A., Sloan D. B., Alden V., Oxelman B. 2012 Phylogenetic relationships of Silene multinervia and Silene section Conoimorpha (Caryophyllaceae). Syst. Bot. 37, 226-237.
253. Reamon-Buttner S. M., Schondelmaier J., Jung C. 1998 AFLP markers tightly linked to the sex locus in Asparagus officinalis L. Mol. Breed. 4, 91-98.
254. Rejon C. R., Jamilena M., Ramos M. G., Parker J. S., Rejon M. R 1994 Cytogenetic and molecular analysis of the multiple sex-chromosome system of Rumex acetosa. Heredity72, 209-215.
255. Renganayaki K., Jessup R. W., Burson B. L., Hussey M. A., Read J. C. 2005 Identification of male-specific AFLP markers in dioecious texas bluegrass. Crop Sci. 45, 2529-2539.
256. Renner S. S., Ricklef R. E. 1995 Dioecy and its correlates in the flowering plants. Am. J. Bot. 82, 596-606.
257. Rice W. R. 1984 Sex chromosomes and the evolution of sexual dimorphism. Evolution38, 735-742.
258. Rice W. R. 1986 On the instability of polygenic sex determination: the effect of sex-specific selection. Evolution40, 633-639.
259. Rice W. R. 1987a Genetic hitchhiking and the evolution of reduced genetic activity of the Y sex chromosome. Genetics116, 161-167.
260. Rice W. R. 1987b Speciation via habitat specialization: the evolution of reproductive isolation as a correlated character. Evol. Ecol. 1, 301-314.
261. Rice W. R. 1987c The accumulation of sexually antagonistic genes as a selective agent promoting the evolution of reduced recombination between primitive sex chromosomes. Evolution41, 911-914.
262. Rice W. R., Gavrilets S., Friberg U. 2008 Sexually antagonistic "zygotic drive" of the sex chromosomes. PLoS Genet. 4, e1000313.
263. Roy R. P., Roy P. M. 1971 Mechanism of sex determination in Coccinia indica. J. Ind. Bot. Soc. 50, 391-400.
264. Ruas C. F., Fairbanks D., Evans R., Stutz H., Andersen W., Ruas P. 1998 Male-specific DNA in the dioecious species Atriplex garrettii (Chenopodiaceae). Am. J. Bot. 85, 162-167.
265. Sackton T. B., Hartl D. L. 2013 Meta-analysis reveals that genes regulated by the Y- chromosome in Drosophila melanogaster are preferentially localized to repressive chromatin. Genome Biol. Evol. 5, 255-266.
266. Sakamoto K., Shimomura K, Komeda Y., Kamada H., Satoh S. 1995 A male-associated DNA sequence in a dioecious plant, Cannabis sativa L. Plant Cell Physiol. 36, 1549-1554.
267. Sakamoto K., Akiyama Y., Fukui K., Kamada H., Satoh S. 1998 Characterization, genome size and morphology of sex chromosomes in hemp (Cannabis sativa L.). Cytologia63, 459-464.
268. Sakamoto K., Ohmido N., Fukui K., Kamada H., Satoh S. 2000 Site-specific accumulation of a LINE-like retrotransposon in a sex chromosome of the dioecious plant Cannabis sativa. Plant Mol. Biol. 44, 723-732.
269. Sakamoto K., Abe T., Matsuyama T., Yoshida S., Ohmido N., Fukui K., Satoh S. 2005 RAPD markers encoding retrotransposable elements are linked to the male sex in Cannabis sativa L. Genome48, 931-936.
270. Samantaray S., Geetha K. A., Hidayath K. P., Maiti S. 2010 Identification of RAPD markers linked to sex determination in guggal [Cammiphora wightii (Arnott.)] Bhandari. Plant Biotechnol. Rep. 4, 95-99.
271. Sanchez-Vilas J., Turner A., Pannell J. R. 2011 Sexual dimorphism in intra- and interspecific competitive ability of the dioecious herb Mercurialis annua. Plant Biol. 13, 218-222.
272. Scotti I., Delph L. F. 2006 Selective trade-offs and sex-chromosome evolution in Silene latifolia. Evolution60, 1793-1800.
273. Seefelder S., Ehrmaier H., Schweizer G., Seigner E. 2000 Male and female genetic linkage map of hops, Humulus lupulus. Plant Breed. 119, 249-255.
274. Segawa M., Kishi S., Tatuno S. 1971 Sex chromosomes of Cycas revoluta. Jap. J. Genet. 46, 33-39.
275. Semerikov V., Lagercrantz U., Tsarouhas V., Ronnberg-Wastljung A., Alstrom-Rapaport C., Lascoux M 2003. Genetic mapping of sex-linked markers inSalix viminalis L. Heredity 91, 293-299.
276. Sharma A. A., Chattopadhyay D. 1991 Sex determination in dioecious species of plants. J. Bot. Taxon. Geobot. 102, 29-55.
277. Sharma A., Zinta G., Rana S., Shirko P. 2010 Molecular identification of sex in Hippophae rhamnoides L. using isozyme and RAPD markers. For. Stud. Chin. 12, 62-66.
278. Shaw A. J., Cox C. J., Goffinet B. 2005 Global patterns of moss diversity: taxonomic and molecular inferences. Taxon55, 337-352.
279. Shiba H., Iwano M., Entani T., Ishimoto K., Shimosato H., Che F. S. et al. 2002 The dominance alleles controlling self incompatibility in Brassica pollen is regulated at RNA level. Plant Cell14, 491-504.
280. Shibata F., Hizume M., Kuroki Y. 1999 Chromosome painting of Y chromosomes and isolation of a Y chromosome-specific repetitive sequence in the dioecious plant Rumex acetosa. Chromosoma108, 266-270.
281. Shibata F., Hizume M., Kuroki Y. 2000 Differentiation and the polymorphic nature of the Y chromosomes revealed by repetitive sequences in the dioecious plant, Rumex acetosa. Chromosome Res. 8, 229-236.
282. Shibu M. P., Ravishankar K. V., Anand L., Ganeshaiah K. N., Shaanker U. 2000 Identification of sex-specification DNA markers in the dioecious tree, nutmeg (Myristica fragrans Houtt). PGR Newslett. 121, 59-61.
283. Shirkot P., Sharma D. R., Mohopatra T. 2002 Molecular identification of sex in Actinidia deliciosa var. deliciosa by RAPD markers. Sci. Hortic. 94, 33-39.
284. Siljak-Yakovlev S., Benmalek S., Cerbah M., Coba de la Pena T., Bounaga N., Brown S. C. and Sarr A. 1996 Chromosomal sex determination and heterochromatin structure in date palm. Sex Plant Reprod. 9, 127-132.
285. Singh R. B., Smith B. W. 1971 The mechanism of sex determination in Rumex acetosella L. Theor. Appl. Genet. 41, 360-364.
286. Siroky L., Lysak M. A., Dolezel J., Kejnovsky E., Vyskot B. 2001 Heterogeneity of rDNA distribution and genome size in Silene spp. Chomosome Res9, 387-393.
287. Skaletsky H., Brown L. G., Kuroda-Kawaguchi T., Minx P. J., Cordum, H. S., Hillier L. et al. 2003 The male-specific region of human Y chromosome is a mosaic of discrete sequence classes. Nature423, 825-837.
288. Smith B. W. 1964 The evolving karyotype of Rumex hastatulus. Evolution18, 93-104.
289. Sondur S. N., Manshardt R. M., Stiles J. I. 1996 A genetic linkage map of papaya based on randomly amplified polymorphic DNA markers. Theor. Appl. Genet. 93, 547-553.
290. Sousa A., Fuchs J., Renner S. S. 2013 Molecular cytogenetics (FISH, GISH) of Coccinia grandis: a ca. 3 myr-old species of cucurbitaceae with the largest Y/autosome divergence in flowering plants. Cytogenet. Genome Res. 139, 107-118.
291. Spada E., Caporali G., Marziani G., Portaluppi P., Restivo F. M., Tassi F. et al. 1998 A genetic map of Asparagus officinalis based on integrated RFLP, RAPD and AFLP molecular markers. Theor. Appl. Genet. 97, 1083.
292. Spielman M., Vinkenoog R., Dickinson H. G. and Scott R. J. 2001 The epigenetic basis of gender in flowering plants and mammals. Trends Genet. 17, 705-711.
293. Spigler R. B., Lewers K. S., Main D. S., Ashman T. L. 2008 Genetic mapping of sex determination in a wild strawberry, Fragaria virginiana, reveals earliest form of sex chromosome. Heredity101, 507-517.
294. Spigler R. B., Lewers K. S., Johnson A. L., Ashman T. L. 2010 Comparative mapping reveals autosomal origin of sex chromosome in octaploid Fragaria virginiana. J. Hered. 101, 107-117.
295. Spigler R. B., Lewers K. S., Ashman T. L. 2011 Genetic architecture of sexual dimorphism in a subdioecious plant with a proto-sex chromosome. Evolution65, 1114-1126.
296. Steflova P., Steflova P., Tokan V., Vogel I., Lexa M., Macas J. et al. 2013 Contrasting patterns of transposable element and satellite distribution on sex chromosomes (XY1Y2) in the dioecious plant Rumex acetosa. Genome Biol. Evol. 5, 769-782.
297. Stehlik I., Blattner F. R. 2004 Sex-specific SCAR markers in the dioecious plant Rumex nivalis (Polygonaceae) and implications for the evolution of sex chromosomes. Theor. Appl. Genet. 108, 238-242.
298. Steinemann S., Steinemann M. 2005 Y chromosome borne to be destroyed. BioEssays27, 1076-1083.
299. Steven J. C., Delph L. F., Brodie E. D. 3rd 2007 Sexual dimorphism in the quantitative-genetic architecture of floral, leaf and allocation traits in Silene latifolia. Evolution61, 42-57.
300. Tanurdzic M., Banks J. A. 2004 Sex-determining mechanisms in land plants. Plant Cell16, 561-571.
301. Telgmann-Rauber A., Jamsari A., Kinney M. S., Pires J. C., Jung C. 2007 Genetic and physical maps around the sex-determining M-locus of the dioecious plant asparagus. Mol. Genet. Genomics278, 221-234.
302. Temsch E. M., Greilhuber J., Krisai R. 2010 Genome size in liverworts. Preslia82, 63-80.
303. Tenaillon M. I., Hufford M. B., Gaut B. S., Ross-Ibarra J. 2011 Genome size and transposable element content as determined by high throughput sequencing in maize and Zea luxurians. Genome Biol. Evol. 3, 219-229.
304. Terauchi R., Kahl G. 1999 Mapping of the Dioscorea tokoro genome: AFLP markers linked to sex. Genome42, 752-762.
305. Testolin R., Cipriani G., Costa G. 1995 Sex segregation ratio and gender expression in the genus Actinidia. Sex Plant Reprod. 8, 129-132.
306. Testolin R., Huang W. G., Messina R., Vecchione A., Cipriani G. 2001 A kiwifruit (Actinidia spp) linkage map based on microsatellites and integrated with AFLP markers. Theor. Appl. Genet. 103, 30-36.
307. Thomas B. C., Pedersen B., Freeling M. 2006 Following tetraploidy in an Arabidopsis ancestor, genes were removed preferentially from one homologue leaving clusters enriched in the sensitive genes. Genome Res. 16, 934-946.
308. Tracey L. P., Koelewijn H. P., Dijk P. J. 2004 Identification of a male- specific AFLP marker in a functionally dioecious fig, Ficus fulva Reinw. ex Bl. (Moraceae). Sex Plant Reprod. 17, 17-22.
309. Turner J. M. A. 2007 Meiotic sex chromosomes inactivation. Development134, 1823-1831.
310. Tuskan G. A., Difazio S., Jansson S., Bohlmann J., Grigoriev I., Hellsten U. et al. 2006 The genome of black cottonwood, Populus trichocarpa (Torr. And Gray). Science313, 1596-1604.
311. Tuskan G. A., Difazio S., Faivre-Rampant P., Gaudet M., Harfouche, A., Jorge V. et al. 2012 The obscure events contributing to the evolution of an incipient sex chromosome in Populus, a retrospective working hypothesis. Tree Genet. Genomes8: 559-571.
312. Uchida W., Matsunaga S., Sugiyama R., Shibata F., Kazama Y., Miyazawa Y. et al. 2002 Distribution of interstitial telomere-like repeats and their adjacent sequences in a dioecious plant, Silene latifolia. Chromosoma111, 313-320.
313. Urasaki N., Tokumoto M., Tarora K., Ban Y., Kayano T., Tanaka H. et al. 2002 A male and hermaphrodite specific RAPD marker for papaya (Carica papaya L.)Theor. Appl. Genet. 104, 281-285.
314. Valenzuela N. 2004 Evolution and maintenance of temperature-dependent sex determination. In Temperature-dependent sex determination in vertebrates (ed. N. Valenzuela and V. Lance), pp. 131-147. Smithsonian Institution Press, Washington, USA.
315. Vallender E. J., Lahn B. T. 2004 How mammalian sex chromosomes acquired their peculiar gene content. BioEssays26, 159-169.
316. Vanburen R., Ming R. 2013 Dynamic transposable element accumulation in the nascent sex chromosomes of papaya. Mob. Genet. Elements3, e23462.
317. van Doorn G. S., Kirkpatrick M. 2010 Transitions between male and female heterogamety caused by sex-antagonistic selection. Genetics186, 629-645.
318. Volz S. M., Renner S. S. 2008 Hybridization, polyploidy, and evolutionary transitions between monoecy and dioecy in Bryonia (Cucurbitaceae). Am. J. Bot. 95, 1297-1306.
319. Vyskot B., Hobza R. 2004 Gender in plants: sex chromosomes are emerging from fog. Trends Genet. 20, 431-438.
320. Vyskot B., Araya A., Veuskens J., Negrutiu I., Mouras A. 1993 DNA methylation of sex chromosomes in a dioecious plant, Melandrium album. Mol. Gen. Genet. 239, 219-224.
321. Wai C. M., Ming R., Moore P. H., Paull R. E., Yu Q. 2010 Development of chromosome-specific cytogenetic markers and merging of broken linkage groups in papaya. Trop. Plant Biol. 3, 171-181.
322. Wang D. W., Li Y., Li Z. Q. 2011 Identification of a male- specific amplified fragment length polymorphism (AFLP) and a sequence characterized amplified region (SCAR) marker in Eucommia ulmoides Oliv. Int. J. Mol. Sci. 12, 857-864.
323. Wang J., Na J. K., Yu Q., Gschwend A. R., Han J., Zeng F. et al. 2012 Sequencing papaya X and Yh chromosomes reveals molecular basis of incipient sex chromosome evolution. Proc. Natl. Acad. Sci. USA109, 13710-13715.
324. Weeks S. C., Benvenuto C., Reed S. K. 2006 When males and hermaphrodites coexist: a review of androdioecy in animals. Integr. Comp. Biol. 46, 449-464.
325. Wellmer F., Riechmann J. L., Alves-Ferreira M., Meyerowitz E. M. 2004 Genome-wide analysis of spatial gene expression in Arabidopsis flowers. Plant Cell16, 1314-1326.
326. Westergaard M. 1958 The mechanism of sex determination in dioecious flowering plants. Adv. Genet. 9, 217-281.
327. Wolf D. E., Satkoski J. A., White K., Rieseberg L. H. 2001 Sex determination in the androdioecious plant Datisca glomerata and its dioecious sister species D. cannabina. Genetics159, 1243-1257.
328. Wu C. I., Xu E. Y. 2003 Sexual antagonism and X-inactivation- the SAX1 hypothesis. Trends Genet. 19, 243-247.
329. Wu X., Wang J., Na J. K., Yu Q., Moore R. C., Zee F. et al. 2010 The origin of the non-recombining region of sex chromosomes in Carica and Vasconcellea. Plant J. 63, 801-810.
330. Wyatt R., Anderson L. E. 1984 Breeding systems in bryophytes. In The experimental biology of bryophytes (ed. A. F. Dyer and J. G. Duckket), pp. 39-64. Academic Press, London, UK.
331. Xu W. J., Wang B.-W., Cui K.-M. 2004 RAPD and SCAR markers linked to sex determination in Eucomma ulmoides Oliv. Euphytica136, 233-238.
332. Yakubov B., Barazani O., Golan-Goldhirsh A. 2005 Combination of SCAR primers and touchdown- PCR for sex identification in Pistacia vera L. Sci. Hortic103, 473-478.
333. Yamato K. T., Ishizaki K., Fujisawa M., Okada S., Nakayama S., Fujishita M. et al. 2007 Gene organization of the liverwort Y chromosome reveals distinct sex chromosome evolution in a haploid system. Proc. Natl. Acad. Sci. USA104, 6472-6477.
334. Yang Z., El Aidi J., Ait-Ali T., Augur C., Teller G., Schoentgen F. et al. 1998 Sex-specific marker and trans-zeatin ribosidase in female annual mercury. Plant Sci. 139, 93-103.
335. Ye D., Installe P., Ciupercescu D., Veuskens J., Wu Y., Salesses G. et al. 1990 Sex determination in the dioecious Melandrium. I. First lessons from androgenic haploids. Sex Plant Rep. 3, 179-186.
336. Yin T., Difazio S. P., Gunter L. E., Zhang X., Sewell M. M., Woolbright S. A. et al. 2008 Genome structure and emerging evidence of an incipient sex chromosome in Populus. Genome Res. 18, 422-430.
337. Younis R. A. A., Ismail O. M., Soliman S. S. 2008 Identification of sex-specific DNA markers for the date palm (Phoenix dactylifera L.) using RAPD and ISSR techniques. Res. J. Agric. Biol. Sci. 4, 278-284.
338. Yu Q., Hou S., Hobza R., Feltus F. A., Wang X., Jin W. et al. 2007 Chromosomal location and gene paucity of the male specific region on papaya Y chromosome. Mol. Genet. Genomics278, 177-185.
339. Yu Q., Hou S., Feltus F. A., Jones M. R., Murray J. E., Veatch O. et al. 2008a Low X/Y divergence in four pairs of papaya sex-linked genes. Plant J. 53, 124-132.
340. Yu Q., Navajas-Perez R., Tong E., Robertson J., Paul H., Moore P. H. et al. 2008b Recent origin of dioecious and gynodioecious Y chromosomes in papaya. Trop. Plant Biol. 1, 49-57.
341. Zhang T., Tan D. Y. 2008 Adaptive significances of sexual system in andromonoecious Capparis spinosa (Capparaceae). J. Syst. Evol. 46, 861-873.
342. Zhang X., Feng B. Q., Zhang D., Altman N., Ma H. 2005 Genome-wide expression profiling and identification of gene activities during early flower development in Arabidopsis. Plant Mol. Biol. 58, 401-419.
343. Zhang Y. H., Di Stilio V. S., Rehman F., Avery A., Mulcahy D. and Kesseli R. 1998 Y chromosome specific markers and the evolution of dioecy in the genus Silene. Genome41, 141-147.
344. Zhou O., Bachtrog D. 2012 Sex-specific adaptation drives early sex chromosome evolution in Drosophila. Science337, 341-345.
345. Zhou Y., Wang X., Zhang X. 2011 Development and application of a SRAP marker for the identification of sex in Buchloe dactyloides. Euphytica181, 261-266.
346. Zluvova J., Janousek B., Negrutiu I., Vyskot B. 2005 Comparison of the X and Y chromosome organisation in Silene latifolia. Genetics170, 1431-1434.
347. Zluvova J., Georgiev S., Janousek B., Charlesworth D., Vyskot B. and Negrutiu I. 2007 Early events in the evolution of the Silene latifolia Y chromosome: male specialization and recombination arrest. Genetics177, 375-386.
348. Zluvova J., Zak J., Janousek B., Vyskot B. (2010) Dioecious Silene latifolia plants show sexual dimorphism in the vegetative stage. BMC Plant Biol. 10, 208.