A definition of sex chromosomes is that they carry sexdetermining genes (i.e.

which control female and male fertility), and that they are heteromorphic
[distinguishable microscopically in the two sexes by size or shape, both from
autosomes and from each other]. When the sexdetermining genes are mapped
to chromosome pairs that are not morphologically different they are called
homomorphic, whereas sex chromosomes ( and ! or " and W) that can be
distinguished by microscopy are classified as heteromorphic (historically called
heterochromosomes or allosomes). When more than two types of sex
chromosomes are found in one indi#idual, they are termed polymorphic. $n some
plants, sex is determined by a simple mendelian genetic system based on the
segregation of one locus, as in s%uirting cucumber (&cballium elaterium), or a
few loci, as in annual mercury ('ercurialis annua).
(heoretical predictions about the existence and organization of these three !
chromosome regions ha#e been confirmed recently by the molecular mapping of
deletion 'utants )uring the past decade, a classical example of plant dioecy has
been re#isited* the li#erwort ('archantia polymorpha). $ndi#idual li#erwort
plantlets (gametophytes) are haploid and possess either an or a !
+hromosome.
&,$-&.&($+/ has a controlling role in animal and plant reproducti#e
de#elopment. 0ne of the best examples of an epigenetic process is the
inacti#ation of one of the two chromosomes in mammalian females, a
phenomenon 1nown as 2arr body formation or lyonization. &pigenetic
mechanisms, especially ).A methylation, are 1nown to ha#e an important role
both in flower setting and in sex organ formation.
/ex expression in plants is often influenced by hormones, as has been well
described in the monoecious models such as the fern +eratopteris richardii and
maize "ea mays. +. richardii is a 30'0/,0405/ fern, and the gender (either
hermaphrodite or male) of its gametophyte is determined by the pheromone
antheridiogen, probably a gibberellin6li1e substance
Recent studies of plant sex chromosomes have characterized successfully the first active genes
that are linked to Silene latifolia sex chromosomes MROS3, SlX !or its homologue "#, SlX$ !or
its homologue "$#, and %%$$& (he lac1 of recombination between proto6 and proto6!
chromosomes e#entually leads to the accumulation of slightly deleterious
mutations and degeneration. ,robably the clearest criterion for characterizing !
chromosome degeneration is a test of whether the ! chromosome is sufficient to
enable the sur#i#al of indi#iduals without at least one copy of 7the ordinary8 sex
chromosome (i.e. the chromosome). 4ecent studies ha#e also re#ealed some
genes on plant ! chromosomes that ha#e no homologues on the chromosome.
(he papaya sex determination system has long been considered to be clear
and simple. A single locus ' with three alleles has a 1ey role in sex
determination ('9, male: ';, hermaphrodite: m, female). 'ny com(ination of t)o
dominant M alleles is em(ryonically lethal& *ecause males are heterozygous and females are
homozygous, sex determination in papaya has (een regarded as the X" type !males are heterogametic,
the male "is dominant#& ,reliminary chromosome analyses show that the papaya sex
chromosomes are homomorphic. RayMing and colleagues demonstrate that degenerative
processes in the non+ recom(ining region of the papaya " chromosome have occurred, indicating that
the first steps of sex chromosome evolution and diversification are under )ay& ,he fact that the
erosion of " chromosome genes in papaya has already (egun is clear from the fact that homozygous
"" em(ryos a(ort -../ days after pollination&
,lant sex chromosomes in dioecious species, although rare, are highly
#ariable and e#olutionarily young, which ma1e them excellent models to study
the structure and e#olution of sex chromosomes. ,lant sex chromosomes can be
grouped into at least four classes illustrating different stages of sex chromosome evolution&