Current Biology

Minireview

Plant Sex Determination

John R. Pannell
Department of Ecology and Evolution, University of Lausanne, 1015 Lausanne, Switzerland
Correspondence: john.pannell@unil.ch
http://dx.doi.org/10.1016/j.cub.2017.01.052

Sex determination is as important for the fitness of plants as it is for animals, but its mechanisms appear to
vary much more among plants than among animals, and the expression of gender in plants differs in impor-
tant respects from that in most animals. In this Minireview, I provide an overview of the broad variety of ways
in which plants determine sex. I suggest that several important peculiarities of plant sex determination can
be understood by recognising that: plants show an alternation of generations between sporophytic and
gametophytic phases (either of which may take control of sex determination); plants are modular in structure
and lack a germ line (allowing for a quantitative expression of gender that is not common in animals); and
separate sexes in plants have ultimately evolved from hermaphroditic ancestors. Most theorising about
sex determination in plants has focused on dioecious species, but we have much to learn from monecious
or hermaphroditic species, where sex is determined at the level of modules, tissues or cells. Because of
the fundamental modularity of plant development and potentially important evolutionary links between
monoecy and dioecy, it may be useful to relax the distinction often made between ‘developmental sex deter-
mination’ (which underpins the development of male versus female flowers in monoecious species) and ‘ge-
netic sex determination’ (which underpins the separation of males and females in dioecious species, often
mediated by a genetic polymorphism and sex chromosomes). I also argue for relaxing the distinction be-
tween sex determination involving a genetic polymorphism and that involving responses to environmental
or hormonal cues, because non-genetic cues might easily be converted into genetic switches.

Introduction and female flowers develop in monoecious plants such as maize

Land plants are remarkably diverse in their sexual systems [1],
but the rich variety of ways in which they express their sexuality
has largely been built on a conserved foundation that determines
whether reproductive tissues produce egg cells or sperm. In this
sense, plants are similar to most other eukaryotes [2]. But when
should the cell cycle end in the ultimate production of eggs
versus sperm, which individuals or flowers should make this
decision, and how should the decision be acted upon in develop-
mental genetic terms? The first of these questions (when?) rec-
ognises that different parts of a plant, at different times, might
be channelled towards being male versus female, and implies
that selection in a given lineage will often optimise a distribution
of decisions among modules in the plant [3,4]. The second ques-
tion (who?) is prompted by the fact that the life cycle of all land
plants involves an alteration of generations between haploid ga-
metophytes and diploid sporophytes, and that one of these
stages might take full or partial control of sex determination
(Figure 1). The third question (how?) refers to the developmental
genetic means by which sex is determined in a given lineage,
with the subtext that there might be some sort of general answer
we might hope to find from a survey among different lineages
[5,6]. In this Minireview, I address these questions by taking an
evolutionary perspective — we should expect sex to be deter-
mined in ways influenced both by a lineage’s phylogenetic his-
tory, and by the action of selection on its populations’ sex ratio
or sex-allocation strategies, ultimately by modifying suitable
gene networks.
For conceptual reasons, I take a broad view of what we mean
by ‘sex determination’. Developmental biologists tend to use the
term ‘sex determination’ when referring to the way in which male
or melon. In contrast, geneticists tend to think of sex determina-
tion as the result of a genetic polymorphism (e.g., in species with
sex chromosomes) that gives rise to the development of males
or females in dioecious species, often initially through male- or
female-sterility mutations. In this genetic sense, the term sex
determination would not apply to monoecious species at all,
because all individuals in a population could be genetically iden-
tical at loci determining sex-organ development.
Although it is useful to remain alert to a possible distinction
in meaning between genetic versus developmental sex determi-
nation, a key point is that the two meanings are not, in fact, as
distinct as they might appear to be. As I discuss below, evolu-
tionary transitions have frequently taken place between dioecy
and monoecy, with the possibility that genes involved in
developmental sex determination in monoecious populations
may play a role in genetic sex determination in dioecious popu-
lations — although this is yet to be established. Moreover, fully
monoecious populations can simultaneously be genetically
dimorphic, with a genetic class of hermaphrodites that empha-
sise their male function (without being exclusively male), and a
second class of hermaphrodites that emphasise their female
function. For these reasons, I blur the distinction between
‘genetic’ and ‘developmental’ definitions of sex determination,
and consider them here together.
The topic of plant sex determination has been reviewed in sub-
stantial detail in recent years [2,5–10]. An important take-home
message is that sex determination in plants is eclectic and varies
a great deal among lineages [5,6]. The contrast with animals is
striking [11]. Whereas much the same gene networks are de-
ployed among highly divergent animals [2,11], the gene networks

Current Biology 27, R191–R197, March 6, 2017 ª 2017 Elsevier Ltd. R191
 Current Biology

Minireview
Figure 1. Relationships between main

Sex chromosomes
clades and grades of land plants.

Control over sex

Heterospory
Illustrated are the key features of variation in:

present

present
whether sex is controlled by the gametophytes,

by
sporophytes or, depending on the lineage, both;
whether hetorospory is present or entirely absent
Non-vascular plants from the group; and whether or not sex is known to
be determined by sex chromosomes. Top image
Bryophytes courtesy of Robert Kipps; centre and bottom im-
ages by the author.
Gametophyte No Yes

Hepatophytes

Vascular
non-seed plants
plant Anthocerophytes phytic phase. However, which phase has
control over the decision, and how the
Lycophytes Gametophyte Yes decision is enacted, varies.
Sporophyte No No
For an overview of sex determination in
Monilophytes land plants, it is convenient to categorise
their diversity into three groups — a non-
Seed plants vascular group, including mosses, liver-
Gymnosperms worts and hornworts (‘bryophytes’ in the
Sporophyte Yes Yes
broad sense); the vascular non-seed pro-
ducing group, including lycophytes and
Angiosperms
monilophytes; and the clade of (vascular)
Current Biology seed plants, including extant gymno-
sperms and flowering plants (angio-
sperms; Figure 1). In non-seed plants, ga-
determining sex even among angiosperms, which have diverged metophytes may be male or female (in ‘dioicous’) species, with
much more recently, appear to be very diverse indeed [5,6]. separate individuals producing either antheridia or archegonia,
Consideration of the nature of plant sexuality, as well as the his- or they may be hermaphroditic (in ‘monoicous’ species), with
tory of transitions between sexual systems, can help to compre- both antheridia and archegonia present in the same individuals.
hend this diversity. Here, sex determination is a matter of the differentiation of cells
In this Minireview, I begin with an overview of a few salient fea- and tissues in different parts of the same individuals. In dioicous
tures of sex determination in the major land-plant lineages species, separate male and female gametophytes germinate
(Figure 1). I then consider what we might infer about plant sex from spores of the same size. Sexual dimorphism nevertheless
determination from the fact that so many plants are hermaphro- can be extreme. In some cases, dwarf male gametophytes
ditic. Functional hermaphroditism in flowering plants takes a grow on larger female gametophytes. In many bryophytes, sex
number of forms, ranging from the case where all flowers are is determined in gametophytes by (U and V) sex chromosomes
‘perfect’, with both male and female functions in each flower, (e.g., [13]).
to ‘monoecy’, where individuals are hermaphroditic but each In ferns and lycophytes, the role and target of sex determina-
flower only develops functional male (stamens) or female organs tion differ between ‘homosporous’ species, in which sporo-
(carpels). Much can be learned about sex determination in plants phytes all produce spores of the same size (as in bryophytes),
by comparing monoecy and dioecy. In both of these sexual sys- and ‘heterosporous’ species, in which sporophytes produce
tems, flowers are either male (staminate) or female (pistillate), but both small ‘microspores’ and larger ‘megaspores’. In some ho-
they differ in whether both sexes can be found in the same mosporous ferns, gametophytes are all functionally hermaphro-
genetic individuals (monoecy) or only in separate genetic individ- ditic, with both antheridia and archegonia. In these cases, sex
uals (dioecy). Finally, I reflect on the implications for our under- determination is a question of cellular and tissue differentiation
standing of sex determination of the fact that plant gender is into different male and female ‘gametangia’. In other homospo-
very often a highly plastic, quantitative trait. rous species, gametophytes may develop a unisexual function
(producing either only antheridia or only archegonia). Here, the
Sex Determination in the Major Land-plant Lineages sex-determination pathway targets separate, individual, game-
The evolution of sexual reproduction constituted a major transi- tophytes. However, the gametophytes differentiate in response
tion in the evolution of life, and occurred well before plants first to environmental cues perceived soon after spore germination
ventured onto land some 470 million years ago. By that time, (Figure 2A), rather than in response to a genetic polymorphism,
the ancestors of land plants already had differentiated male as in some mosses mentioned above. Indeed, to my knowledge,
and female gametes (spermatozoids and eggs), and different sex chromosomes have not been described for any ferns or
sex organs to produce them (archegonia and antheridia). The lycophytes [6].
first land plants had also already evolved an alteration of haploid In ‘heterosporous’ ferns and lycophytes, sex determination
and diploid generations (the gametophytes and sporophytes) acts through the size of the spore from which the gametophytes
[12]. In all land plants, the differentiation of sex organs into germinate. The gametophytes do not differ genetically, and the
egg- or sperm-producing tissues occurs in the haploid gameto- sporophyte controls the sex of its gametophytes by regulating

R192 Current Biology 27, R191–R197, March 6, 2017

Current Biology

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A the spore-producing ‘sporangia’ to produce either small ‘micro-

HERs + – HERs spores’, which develop into male (micro-) gametophytes, or
larger ‘megaspores’, which develop into female (mega-) gameto-
TRAs MAN1 NOT1 TRAs phytes. Heterospory has evolved several times in vascular land
plants, in ferns and lycophytes, as well as independently in the
NOT1 FEM1 FEM1 MAN1 lineage that gave rise to seed plants [14].
In angiosperms (flowering plants) and gymnosperms, the
Male Exogenous cue: Female gametophytic phase has become even more highly modified
gametophyte antheridiogen gametophyte and reduced, and differentiation between the male and female
structures takes place in sporophytic structures in which the ga-
metophytes form. In extant seed plants, the role of sex determi-
nation is thus passed from the gametophytes entirely to their
diploid sporophyte ‘hosts’. Seed-plant sporophytes may ulti-
mately be functionally hermaphroditic, producing both pollen
(modified and highly reduced male gametophytes) and ovules
B (which contain modified female gametophytes), or they may be
OGI XY XX
male or female, producing only pollen or ovules. In contrast to
MeGI MeGI
non-seed plants, seed-plant gametophytes are always either
male or female, and it is the sporophyte that determines their
sex. The developmental machinery of sex determination in
Stamen Stamen
development
seed plants therefore governs which gametophytes a given
development
diploid sporophyte individual produces, where on the plant
they will occur, and in which proportions (i.e., the sex allocation;
see below).

Implications of Widespread Hermaphroditism for Sex

Determination in Plants
Male flower Female flower
The great majority of plants are hermaphroditic angiosperms
[1,15,16]. Most of these have ‘perfect flowers’, with a remarkably
C conserved arrangement of their four floral whorls, the sepals,
ACS11 ON OFF ACS11 petals, stamens, and the central carpels. The genetics of floral
development, which also implies sex differentiation in specific
WIP1 ASC-7 WIP1 ASC-7 tissues, is now reasonably well characterised in terms of the
ABCE model [17,18], which envisages organ identity as the
Carpels Stamens Carpels Stamens outcome of (partially overlapping) expression of genes in four
gene classes. In particular, sex is determined by expression of
Endogenous cue:
ethylene B- and/or C-class genes that initiate the development of sta-
mens or carpels [19]. Although the action of these gene classes
for organ differentiation in flowers has been investigated in detail
in only a handful of species, it appears to apply quite generally
across flowering plants, and probably evolved with the origin
Female flower Male flower
Current Biology of flowers [19]. Given the highly conserved role of a number of
genes in sex determination in animals [2], one might have ex-
Figure 2. Three contrasting sex-determining pathways in land pected the ABCE genes to play a key role in determining sex in
plants. dioecious species. However, although they do play a key role
Black and grey text and symbols indicate genes and functions switched on in some lineages, they are rarely the genes that decide whether
and off, respectively. Arrows and ‘flat-ended’ arrows indicate promotion or
a plant becomes male or female [5,6].
suppression of activity, respectively. (A) Environmental determination of sex in
the homosporous fern Ceratopteris richardii. Sex determination depends on The wide diversity of genetic mechanisms for sex determina-
the outcome of two antagonistically interacting genes, FEM1 and TRA: in the tion in dioecious plants is attributable to the fact that separate
absence of an exogenous signal from the hormone antheridiogen, the gene sexes have evolved repeatedly in different lineages, often
TRA is expressed; this both elicits the development of a female (or her-
maphrodite) gametophyte, and suppresses the expression of FEM1 through recently [15,20], and that genetic switches involved in sex
an intermediary MAN1. In the presence of exogenous antheridiogen, the gene
HER is expressed; HER suppresses the expression of TRA (and, in turn, of
MAN1), allowing the expression of FEM1. FEM1 promotes both the develop-
ment of a male gametophyte and the expression of the gene NOT1, which in The model invokes three interacting loci, with gene ASC11 suppressing the
turn suppresses TRA. Images courtesy of Jody Banks. (B) Chromosomal carpel suppressor WIP1, and WIP1 also suppressing the stamen suppressor
determination of sex in persimmons. In XY individuals (males), expression of ACS-7, allowing flowers to develop carpels and to be fully female; expression
OGI, a small RNA, targets and suppresses a second autosomal gene MeGI, a of WIP1 thus both prevents the production of carpels and suppresses ACS-7,
transcription factor that suppresses stamen development. In XX individuals, allowing flowers to develop stamens and thus to be fully male. An upstream
which lack OGI, MeGI is expressed, suppressing stamen development. Im- regulatory cue (perhaps the hormone ethylene) turns ASC11 on or off in
ages courtesy of Jeff Pippen. (C) Sex determination in monoecious melons. different parts of the plant. Images courtesy of Marcel Dorken.

Current Biology 27, R191–R197, March 6, 2017 R193


A Figure 3. The quantitative (and potentially
1.0
1.0
Relative rank
0.5 0.5
0 0
0 0.5 1.0
Proportion of female flowers in inflorescence
B Cossard (centre left) and Xinji Li (right).
Female (XX) ‘Leaky’ female (XX)
determination have thus evolved many times independently
[5,21]. But generalisation is nonetheless possible if we consider
how transitions to dioecy have likely occurred. First, dioecy must
evolve through the positive selection of both male and female
sterility mutations in what are destined to become the sex-deter-
mination genes. Theory predicts that male-sterility mutations are
most likely to spread in hermaphrodite populations first, largely
because they prevent self-fertilizing, and thus from producing
progeny that might suffer inbreeding depression, but also
because it is difficult for female-sterility mutations to spread
in partially selfing populations [22,23]. The ‘gynodioecious’ pop-
ulation that results may then evolve towards full dioecy, for
example, in response to selection for sexual specialisation,
either with the subsequent spread of a female-sterility mutation,
or through the gradual shift in sex allocation of non-male-sterile
hermaphrodites towards greater maleness (the production of
fewer seeds and more male flowers or pollen) [23,24].
The two likely alternative paths from gynodioecy to dioecy
have important implications for how sex might be determined.
In the case of the spread of a female-sterility mutation in a gyno-
dioecious population, the gene affected must become tightly
linked to the male-sterility mutation on a single (sex) chromo-
some, because otherwise segregation or recombination could
create genotypes that carry and express both male and female
sterility at the same time [22]. Alternatively, the female-sterility
mutation could show epistatic expression, masculinising only
those individuals not expressing the male-sterility mutation. If
R194 Current Biology 27, R191–R197, March 6, 2017
Current Biology
Minireview
plastic) nature of sex determination in
plants.
(A) Plots of the proportion of female flowers in two
dioecious species, taken from the work of D. Lloyd
(republished with permission from John Wiley and
Sons, Inc., [50]), who first emphasized the impor-
tance of quantitative gender in plants. In both
plots, the distribution of gender is plotted as the
relative rank gender of individuals sampled in the
population, with closed and open circles repre-
senting male and female plants, respectively. Left
panel: Gingidia montana; right panel: Leptinella
dendyi. (B) Chromosomal sex determination in the
plant Mercurialis annua, showing an XX female
(left), an XY males (centre), and an XY male with
‘leaky’ gender expression, i.e., a male producing
0 0.5 1.0 both male and female flowers and fruits. Blue ar-
rows indicate male (staminate) flowers; red arrows
indicate female (pistillate) flowers. Images cour-
tesy of John Baker (left and centre right), Guillaume
male sterility is recessive, as is most
probable for a loss-of-function mutation
[22,25], females should be homozygous
(and thus XX in the case of a sex chromo-
some), and hermaphrodites or males
should be heterozygous (and thus XY,
with female sterility or male enhancement
Male (XY) ‘Leaky’ male (XY) genes being located on the Y).
Little is known about the genes that
Current Biology
cause male and female sterility in dioe-
cious plants, but it seems that genetic
linkage between them on a sex chromo-
some is common [26], as exemplified by recent work on white
campion (Silene latifolia) [27], wild strawberry (Fragaria virginiana)
[28] and broadleaf arrowhead (Sagittaria latifolia) [29]. Papaya
(Carica papaya) also has an XY sex-chromosome system, but
a modified Yh chromosome is also present, and that fails to sup-
press the individual’s female function, allowing hermaphrodites
to develop, and suggesting that the normal Y in papaya males
indeed harbours a female suppressor [30]. Modified Y chromo-
somes are similarly known in Silene latifolia [26] and Vitis vinifera
[31]. In the annual mercury (Mercurialis annua), females have no
Y chromosome, and have acquired a male function in some pop-
ulations [32] (Figure 3B), suggesting either that male-sterility
mutations on an X chromosome have incomplete expression in
these individuals, or that there are loci elsewhere in the genome
that can partially restore male function in females.
The two-locus model for the evolution of sex determination
and dioecy makes intuitive sense and has received some sup-
port [8], but sex is determined in many species by a single switch
gene [33]. In Caucasian persimmon (Diospyros lotus), for
example, the anther fertility is determined by transcription of a
Y-linked sequence encoding a small RNA that targets and sup-
presses expression of an unlinked autosomal feminizing gene
(Figure 2B) [34]. In other words, although two loci are involved
at the top of the sex-determination cascade, they are unlinked
to one another, and only one of them now acts as the switch
gene in the gene network. It is not yet known how such develop-
mental gene networks have been assembled during the
Current Biology
Minireview
evolution of dioecy. For example, did they evolve through the
initial spread of (linked) male- and female-sterility mutations, as
supposed by theory [22]? Or might they have evolved through
the spread of a single sex-determining gene distinguishing be-
tween females and hermaphrodites, say, followed by the fixation
of mutations expressed only in hermaphrodites that reduce the
female allocation of the hermaphrodites?
The plausibility of single-locus sex determination in some spe-
cies is also suggested by the ontogeny of male and female ste-
rility. In their review, Diggle et al. [5] noted that stamen abortion in
a given species tends to occur at approximately the same time
as carpel abortion. This pattern is not what we would immedi-
ately expect if sexual organ abortion results from the expression
of independent male- and female-sterility mutations, unless it re-
flects a wide distribution of times since the evolution of dioecy in
the respective lineages. Alternatively, the pattern hints at the
possibility that a single underlying switch acts within a gene
network to suppress one or the other sexual function at about
the same time of floral differentiation [5]. Such a scenario has
been elucidated in monoecious melons [35,36], although again
it is by no means clear how it evolved.
In melons, and in maize, the sexual identity of flowers appears
to result from the expression of genes in different networks that
are responsive to positional information mediated by hormones.
In melons, gene expression at one locus causes suppression of
carpel development (female function), whereas expression at a
different locus suppresses stamen development (the male func-
tion). Expression of an upstream gene that is thought to be sen-
sitive to local ethylene levels determines which of those two
genes is switched on (Figure 2C) [35]. In maize, another monoe-
cious species, different mutant classes have been identified that
suppress stamen or carpel development in female (ears) or male
inflorescences (tassels), respectively [37]. Just as male and fe-
male flowers in melons tend to develop in different parts of the
plant, ears and tassels in maize develop in leaf axils and at the
plant apex, respectively; the gender distribution within the plant
is responsive to gradients in plant hormone levels [38], and con-
forms to adaptive expectations for a wind-pollinated plant [39].
The Quantitative and Plastic Nature of Plant Sex
Determination
Plant gender is fundamentally a quantitative trait [40]. Whereas
individuals in most gonochoristic animals are either fully male
or female, gender in plants is often better viewed on a quantita-
tive scale, measured in terms of the relative allocation to each
sexual function, or the proportion of genes transmitted through
sperm versus eggs (Figure 3A) [41]. Many dioecious species,
including those with well-developed sex chromosomes, show
‘leaky’ or ‘inconstant’ gender expression, with males and fe-
males producing flowers of the other gender. Far from being
an aberrant feature of plant reproduction, such phenotypes are
typically fully functional for both sexes (e.g., Figure 3B). The
quantitative distribution of gender in a population is under strong
selection and can evolve rapidly when circumstances change —
for example, when males are lost from a population [42]. Such
shifts in sex allocation can occur at any level in the modular hier-
archy of plant development, from the number of pollen grains
produced in anthers, to the number of floral or inflorescence
primordia that develop with male versus female functions [40].
Such variation allows for multiple pathways to sex differentiation
and potentially sex determination (Figure 3).
While the distribution of quantitative gender in a population
may have a strong genetic component, individuals can also
modify their sex expression in response to context-dependent
opportunities or costs [40]. Plastic sex expression applies not
only to the quantitative nature of sex allocation in functional her-
maphrodites [43], but, more rarely, to complete sex change [44].
In jack-in-the-pulpit (Arisaema species), young plants are fully
male, whereas, in later seasons, they become fully female or her-
maphroditic [45]. Such ‘sexual diphasy’ is expected to evolve
both if the marginal benefits of reproducing as male versus fe-
male change with plant size [46], and if there are additional ben-
efits to sexual specialisation at a particular point in time (e.g., if
fitness is compromised by interference between simultaneous
male and female functions) [47]. Plasticity in sex allocation, and
sexual diphasy specifically, indicate that sex may be determined
not by a genetic dimorphism, but by the differential expression of
genes shared by all individuals in a population in response to
physiological and environmental cues.
Sex in jack-in-the-pulpit is likely to be responsive to intrinsic
physiological signals reporting the resource status of the plant,
but also to exogenous cues. Such responses have been docu-
mented for many angiosperm species, although angiosperms
do not seem to have evolved fully environmental sex determina-
tion (ESD) that might parallel, for example, temperature-depen-
dent sex determination in some reptiles. In contrast, some homo-
sporous ferns do show true ESD [6,48,49]. When the haploid
spores of the fern Ceratopteris richardii germinate, for example,
they develop into either male or female gametophytes, depend-
ing on the presence of potential mates (Figure 2A). In the absence
of mates, gametophytes produce both antheridia and arche-
gonia. They also release into the water around them the hormone
antheridiogen, which causes nearby gametophytes to develop as
miniature males. Some of the underlying genes involved in the
switch between hermaphroditic and male pathways have been
identified [6,48]. Mutations at these loci render gametophytes
insensitive to antheridiogen, and their sex is thus determined
genetically by the mutations they carry (Figure 2A) [6,48]. In prin-
ciple, the spread of such mutations in a natural population could
conceivably convert ESD into genetic sex determination (GSD). In
monoecious melons cited above, or in maize, in which the sex of
individual flowers or inflorescences depends on their location in
the plant, genetic manipulation of mutants can render a popula-
tion effectively dioecious in similar ways [35,36], that is, experi-
mental manipulation can convert a type of hormone-sensitive,
tissue-specific ESD into GSD. Such examples show that ESD
and GST may be readily interconverted.
Conclusions
Much has been learned about how sex is determined in a handful
of plant species. Although detailed analysis of a wider taxonomic
sample will no doubt be revealing, it is already becoming clear
that different gene networks probably determine sex in different
monoecious and dioecious lineages, and that generalisations
are thus unlikely to emerge in terms of the classes of genes
involved. However, the very diversity of sex-determination
mechanisms in plants reflects the important fact that separate
sexes tend to have evolved independently in different lineages,
Current Biology 27, R191–R197, March 6, 2017 R195

and much more recently than in most animal lineages. This diver-
sity should not surprise us.
But are there any general patterns in how gene networks come
to be assembled that involve the co-ordination of both male- and
female-sterility mutations, often in different parts of the plant or-
ganism? The two-locus model for the evolution of sex chromo-
somes and sex determination is satisfying in its potential gener-
ality and logical coherence. In contrast, we have no similarly
general proposition for how single-locus sex switches become
embedded in necessarily complex networks. For instance, do
dioecious species employ the same developmental pathways
as the monoecious species from which they are derived? Ad-
dressing these questions should be a major goal for future
research, for example, through a comparative analysis of gene
networks in related species that express their gender differently.
The questions of when and where on a plant which sex is ex-
pressed has long been accessible through analysis of competing
allocation strategies in an evolutionary framework. The theoret-
ical underpinnings for such an understanding are well-estab-
lished [3], and observed patterns of sex expression and sex allo-
cation often conform nicely to expectations – though many
puzzles remain. Because sex determination directly affects an
important currency of fitness (reproductive success), it is not sur-
prising that its responses to selection can be rapid and dramatic.
An interesting question that deserves further attention here
concerns shifts between plasticity and genetic polymorphism.
How often plasticity in sex expression acts as a capacitor for
the evolution of genetically separate sexes is not known, but
the facility with which these modes of sex determination can
be interconverted experimentally suggests that they do not
reflect fundamental evolutionary constraints. Nonetheless, it
remains puzzling as to why a chromosomal polymorphism that
determines sex in the gametophytes of bryophytes has never re-
placed environmental sex determination in ferns or lycophytes. It
is tempting to think that the distinction is linked to the ephemeral
nature of fern but not bryophyte gametophytes [6], but why there
should be such a link is not obvious.
ACKNOWLEDGEMENTS
I thank Jörn Gerchen, Paris Veltsos, Wen-Juan Ma and three anonymous re-
viewers for comments on the manuscript, Deborah Charlesworth and mem-
bers of my lab group for valuable discussions, and the University of Lausanne
and the Swiss National Science Foundation for funding.
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