Bryophytes
Monoecious bryophytes often exhibit temporal variations in sex distribution, influenced by
environmental factors such as density and nutrient concentration. Additionally, hormonal treatments
have been found to influence sex expression in some species, with auxin and gibberellin stimulating
maleness and cytokinins stimulating femaleness in certain mosses. Similarly, growth-related
substances like gibberellin and ethrel promote antheridial formation, while auxin and glycocel
enhance archegonial development in liverworts.
Overall, bryophytes display a complex interplay between genetic and environmental factors in sex
determination and expression. Despite limited experimental data, it's evident that environmental
cues play a significant role in shaping the sexual characteristics and reproductive strategies of these
ancient plants.
Pteridophytes
Pteridophytes, comprising ferns and fern allies, exhibit two main classifications: homosporous and
heterosporous. In their life cycle, a haploid gametophyte alternates with a diploid sporophyte.
Homosporous species, predominant among pteridophytes, produce spores of one size, while
heterosporous species produce spores of two size classes. Heterospory is limited to specific groups
like Isoetaceae, Selaginellaceae, aquatic ferns, and certain terrestrial ferns. In heterosporous species,
microspores and megaspores develop into haploid male and female gametophytes, respectively,
contrasting with homosporous species where spores of the same size may develop into male, female,
or hermaphroditic gametophytes. Pteridophyte gametophytes are often termed hermaphrodites,
though monoecious might be more accurate due to separate locations of antheridia and archegonia.
Among homosporous pteridophytes, sex expression is variable and influenced by factors like
gametophyte age, size, and antheridiogen, a gibberellin-like hormone secreted by neighboring
gametophytes. Antheridiogens facilitate male development in the presence of rapid gametophyte
growth.
Studies on Ceratopteris richardii, a homosporous fern, elucidate genetic and environmental factors
affecting sex determination. Antheridiogens play a crucial role in initiating and maintaining male
expression, promoting inter gametophytic matings, and controlling genetic variation.
Approximately 10% of pteridophyte species are agamosporous, reproducing without meiotic
reduction via apogamy or apomixis. Some fern families have eliminated the sporophyte stage,
reproducing vegetatively via gemmae.
The dynamics of sex expression, antheridiogen production, and gametophyte mating strategies
among pteridophytes underscore their adaptability and complex reproductive biology, offering
insights into plant evolution and diversity.
Prepared by Sourav R Mohapatra, Asst. Professor, Dept. of FBT, College of Forestry
SEX EXPRESSION
Introduction:
Flowers can be bisexual (monoclinous), have unisexual flowers of both sexes on the same
individual (monoecy), or have unisexual flowers on different individuals (dioecy). Monoecy
is relatively common in the plant kingdom, with estimates suggesting a frequency slightly
higher than dioecy.
Floral diversity
Individual plants can exhibit different sex types, including monoclinous individuals
(all H), monoecious (M + F), gynomonoecious (H + F), andromonoecious (H + M), and
trimonoecious (H + F + M). Unisexual individuals exist as either male (M) or female (F).
Different combinations of these sex types give rise to various sexual systems, such as dioecy
(male and female), gynodioecy (female and monoclinous), androdioecy (male and
monoclinous), trioecy (male, female, and monoclinous), unisexual females with
agamospermy, and monocliny (monoclinous only). The botanical terminology of floral sex
originates from Linnaeus' systema sexualis, which used the metaphor of a marital bed.
Linnaeus classified flowers as monoclinous (one bed) or diclinous plants being either
monoecious or dioecious. The term "polygamous" was used for plants with mixed flowers,
but it was later replaced with terms like gynomonoecy, andromonoecy, and trimonoecy.
However, "polygamy" still serves as a catch-all term for any mixture of monoclinous and
diclinous flowers.
Quantitative gender Theory
David Lloyd's seminal papers, notably "Sexual strategies in plants III: A quantitative method
for describing the gender of plants," introduced the concept of "quantitative gender" to
describe this phenomenon. Lloyd defined gender as the quantitative estimation of an
individual plant's maleness or femaleness, based on the proportions of its genes transmitted
through pollen or ovules. Gender is represented by a formula where Gi represents the
femaleness gender, and Ai (maleness gender) is 1 − Gi. This concept applies specifically to
cosex systems, including monoecious, polygamous, or pleogamous plants. In strict dioecy,
gender coefficients are fixed at 1 and 0.
Two gene model (Monecy sex expression):
Sex expression of monoecy from monocliny (hermaphrodite) typically requires two
mutations: a feminizing mutation (F) to produce female flowers and a masculinizing mutation
(M) to produce male flowers. These mutations may involve separate loci (genetically FFMM)
or a single gene with two genetic changes in transcriptional regulation. For instance, a stamen
developmental gene (M) could undergo a dominant mutation, gaining function to be
expressed throughout the floral meristem, thereby masculinizing the flowers. Subsequently,
this gene could acquire a cis-regulatory element in the promoter allowing susceptible to
repression, thus becoming the feminizing factor F. Importantly, these mutations typically
involve gain of function rather than loss of function, as monoecious plants have fully
functional sexes.
Prepared by Sourav R Mohapatra, Asst. Professor, Dept. of FBT, College of Forestry
Figure 1. Sex expression in Monoecy
Single-gene Model (Expression of dioecy):
Dioecy is controlled by a single gene M, with a strong allele (superscript plus; M+), and null
allele (superscript minus; M−). It will periodically give XY sex-determination system. The
simplest explanation for the expression of dioecy involves mutations in a single gene,
particularly one involved in sexual determination. In the scenario described, genes controlling
feminization (F) and masculinization (M) are present in the monoecious state, denoted as
FFMM. This genetic setup allows for the development of both male and female flowers
within the same plant. The transition to dioecy typically requires two mutations in one of
these genes. For instance, in the XY sex-determination system (as observed in Populus
tremula), a strong allele of gene M (M+) leads to the development of male flowers (FFMM+),
while a weak or null allele of M (M-) results in female flowers (FFM-M-). This genetic
configuration segregates in an XY pattern, where males are heterogametic.
Figure 1. Sex expression in Dioecy
Prepared by Sourav R Mohapatra, Asst. Professor, Dept. of FBT, College of Forestry
