Flower-type diversity and its evolutionary history
Background:
Around 2000 years ago, Egyptians were already understanding the basics of plant reproduction by
crossbreeding male and female date palm plants to produce fruits. However, in-depth studies on flower
pollination began much later, around the 17th and 18th centuries, with naturalists like Christian Konrad
Sprengel, Joseph Ko¨lreuter, and Thomas Knight. These early efforts paved the way for Charles
Darwin's extensive observations and experiments on plant reproduction in the 19th century. In the
1930s-1950s, the neo-Darwinian synthesis sparked interest in the genetics of plant reproductive
systems. Scientists like Ronald A. Fisher and J. B. S. Haldane explored the inheritance of sexual traits
in plants. Fisher, in particular, analyzed conditions influencing the spread of a gene leading to self-
fertilization, applying population genetic principles to mating-system evolution. By the 1970s, there was
a shift from a group-selection perspective to individual selection models. During this time, researchers
started using genetic markers, like allozyme polymorphisms, to directly measure mating parameters in
plant populations. This allowed for more accurate estimates of outcrossing rates, inbreeding
coefficients, and paternity.Until the 1970s, most research in plant reproductive biology was descriptive.
However, a new theoretical framework emerged in the 1970s through the work of scientists like David
Lloyd, Deborah and Brian Charlesworth, and Eric Charnov. This marked the beginning of a new era in
plant reproductive biology, focusing on concepts like fertility, gender equality, population genetic, and
phenotypic selection model.
Evolutionary transition
Evolutionary transitions are functional changes to adaptive traits that spread to replace ancestral
conditions because they increase fitness.
Evolution of Selfing flowers:
Two main explanations for the evolution of selfing are: (i) advantage in situations of scarce pollinators
or mates (reproductive assurance), and (ii) genetic transmission advantage through pollen, where
selfing variants act as both maternal and paternal parents (automatic selection). Biogeographical
evidence supports the idea that selfing populations occupy areas with reduced pollinator densities, as
predicted by the reproductive assurance hypothesis. However, experimental evidence for this
hypothesis is limited. Additionally, there is relatively less research on the automatic selection
hypothesis.
(iii) Heterostyly and Transition to Selfing:
Heterostylous species have two or three distinct mating morphs with reciprocal arrangements of sexual
organs, promoting pollinator-mediated disassortative mating. Some heterostylous species transition
from obligate outcrossing to predominant selfing. This shift is often associated with the development of
homostyles, where anthers and stigmas are in close contact, allowing for self-pollination.
(iv) Genetic Mechanisms in Turnera and Eichhornia (Case study):
In Turnera, homostyly arises through recombination between linked loci governing the
style-length/anther-height polymorphism. In Eichhornia, homostylous selfing forms arise initially through
major gene changes to stamen position, followed by polygenic modifiers reducing flower size.
Selfing homostyles in both Turnera and Eichhornia tend to occupy geographical range margins,
supporting the concept of reproductive assurance. Colonization of Caribbean islands by homostyles is
observed due to the capacity of single individuals to establish colonies.
Prepared by Sourav R Mohapatra, Asst. Professor, Dept. of FBT, College of Forestry
Persistence of Selfing Lineages:
The question of whether selfing lineages persist and contribute to evolutionary patterns (phyletic
evolution) is unresolved. However, the degree of selfing matters; highly selfing species with specific
traits (selfing syndrome) may face limitations in re-evolving outcrossing. However, selfing species with
moderate outcrossing rates might have the potential to return to higher outcrossing levels if there is
sufficient genetic variation for traits affecting pollinator visitation within populations.
Evolution of Dioecy:
Theories:
1. Darwin's Puzzle:
Darwin struggled to understand why some angiosperm species would abandon hermaphroditism in
favor of dioecy. In plants, where mobility is limited, the immediate advantage of hermaphroditism is not
clear since losing pollen vectors could compromise individual fitness.
2. Genetic Transmission and Sterility Mutations:
Dioecy involves separate male and female individuals, reducing genetic transmission for unisexual
compared to hermaphrodites. The evolution of dioecy requires the spread of sterility mutations, which,
in most circumstances, would be deleterious to fitness.
3. Inbreeding Avoidance:
The most widely accepted hypothesis for the function of dioecy is inbreeding avoidance, preventing
self-fertilization. Gender dimorphism (Dioecy) commonly evolves from self-compatible ancestors rather
than self-incompatible ones, supporting the anti-selfing hypothesis.
Evidence from Saggitaria latifolia:
The study provides evidence that selfing rates (the proportion of seeds produced through self-
fertilization) and levels of inbreeding depression (reduction in fitness due to mating between close
relatives) in monoecious populations of S. latifolia are observed to exceed theoretical values. The
higher-than-expected selfing rates and inbreeding depression suggest that self-fertilization within
monoecious populations might be more prevalent and impactful than predicted by theoretical models.
The findings indicate that if inbreeding depression is strong, the conditions might favor the spread of
unisexual variants, potentially leading to the establishment and persistence of dioecious populations.
Occurrence and Evolution of Dioecy:
1. Frequency and Routes:
Dioecy is not very common in flowering plants (approximately 6–7% of species) but occurs in close to
half of all angiosperm families, having evolved on at least 100 occasions. Two main evolutionary routes
to dioecy: the "gynodioecy pathway" (involving populations with females and hermaphrodites) and the
"monoecy pathway" (involving selection on sex allocation in monoecious ancestral populations). The
transition from monoecy (individuals with both female and male flowers) to dioecy has been relatively
neglected in research. Recent work on Sagittaria suggests that dioecy may evolve from monoecy via
the gynodioecy pathway.
2. Traits Associated with Dioecy:
Prepared by Sourav R Mohapatra, Asst. Professor, Dept. of FBT, College of Forestry
Dioecy is commonly associated with certain life history and reproductive traits. It occurs most commonly
in long-lived species and is rare in annuals. Dioecy is associated with several traits, including the
abiotic pollination, small inconspicuous white or green flowers, and fleshy fruits.
Prepared by Sourav R Mohapatra, Asst. Professor, Dept. of FBT, College of Forestry
