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on
Subject: Genetics
M.Sc. Botany
(2023 – 25)
Department of Botany
1.1.1 Self-pollination
Technically, Self-pollination is the phenomenon of a flower being pollinated by pollen from a
flower of the same plant or the pollen of a different flower but from the same plant. A very
high rate of Self-pollination has important genetic and evolutionary impacts those are
responsible for influencing the genetic structure of a population, gene diversity, evolutionary
rates, effective population size, and patterns of evolutionary diversity(Barrett Spencer C.
H.;2010). The molecular mechanism concerned with corolla size reduction in the selfing
species of Capsella has been explored, where the selfing species of Capsella rubella has
flowers with an 85% reduction in petal size, it’s outcrossing species i.e. Capsella grandiflora
bears corolla of normal size. Researchers informed that a region in the intron of the STERILE
APETALA (SAP) gene, that encodes an F-box protein ( Byzova MV, Franken J, Aarts
MGM, de Almeida-Engler J, Engler G, et al.; 1999), was responsible for a 25% reduction
in petal area of C. rubella as compared to C. grandiflora. Also, it has been reported that in
Eichhornia, homostylous(having the same size of stylous) forms that undergo self-pollination
arise because of the major gene changes of stamen position along with further polygenic
changes that resulted in a reduction of flower size (Barrett et al. 2009). Theoretical models
prepared on the co-evolution of selfing and inbreeding depression suggested a general pattern
(Lande & Schemske 1985), this further resulted in significant experimental work on the
connection between selfing rates and the life-history dynamics revolving around inbreeding
depression.
1.1.2 Cross-pollination
Cross-pollination is the phenomenon of transfer of pollen from anther of one plant to the
stigma part of the pistil of another flower. Cross-pollination serves various advantages to
plants and one of the prominent advantages of the same is an increase in genetic diversity
thus providing selection during challenging situations - the basic rule of genetics.(Fattorini,
R., & Glover, B. J.;2020).The higher genetic diversity also results in a great increase in
phenotypic variation promoting a greater diversity of visitors (Barbour et al. 2020). The
most widely accepted hypothesis given by researchers, for the usefulness of dioecy(unisexual
flowers), is that it provides a brilliant escape from the inbreeding depression by preventing
self-fertilization. Theoretical models suggested by researchers identify three major sets of
factors that are important while considering the evolution of dioecy(unisexuality): selfing and
inbreeding depression, the optimal availability of resources to female and male plants, and
the genetic control of sex determination (Charlesworth 1999). Self-incompatibility (SI) is an
adaptive strategy in plants that increases genetic variability by promoting cross-pollination.
1.3 Self-incompatibility
Self-incompatibility(SI) is a genetic mechanism where a flower rejects genetically
related(self) pollen considering it incompatible and this is how we derived this word. This
increases genetic diversity in plants by promoting cross-pollination instead of
self-pollination, ultimately reducing inbreeding depression. In self-incompatible plants,
pollination is expected to reduce seed production if self-pollen comes to stigma, it also causes
damage to outcrossed pollen tube growth, increases chances of fruit abortion, and also
reduces chances of pollination success(reviewed in Snow et al., 1996). Gender dimorphism
commonly evolves from self-compatible rather than self-incompatible ancestors, a pattern
consistent with the anti-selfing hypothesis.
The SI response of pollen is concerned either with the genotype of the diploid parent plants
that produce pollen (sporophytic self-incompatibility, SSI) or the genotype of the haploid
microspores (gametophytic self-incompatibility, GSI). In either of these cases, the
incompatibility mechanism is under the influence of genes located in the multi-allelic S
loci(Chakraborty, S., Dutta, S. & Das, M.; 2023).In the case of GSI, self, and
non-self-pollens are initially allowed to germinate. After that only non-self pollen is allowed
to enter the ovary for fertilization, and the growth of self-pollen is stopped in style. In SSI, a
receptor kinase-based interaction between S LOCUS CYSTEINE RICH (SCR) and S
LOCUS RECEPTOR KINASE (SRK) genes influences the incompatibility response.
According to research studies, at the molecular level, SI is controlled by two tightly linked
genes present in the S locus (Muñoz-Sanz et al. 2020).SSI occurs when even one of the S
proteins present in pollen matches with at least one of the two S proteins present in the pistil.
In the case of GSI, the SI mechanism is concerned with the genotype of the haploid
microspores (Silva and Goring 2001).GSI occurs when the S haplotypes of pollen match
with at least one of the S haplotypes in a heterozygous pistil.
The female factor of the S locus gene is the S LOCUS RECEPTOR KINASE (SRK). Similar
to the SRK component of the pistil, the male factor is the S LOCUS CYSTEINE RICH /S
LOCUS PROTEIN 11 (SCR/ SP11).SCR and SRK proteins ultimately influence and produce
SI responses. In the last 25 years, the molecular characterization of the S-locus in species of
the various economical families including Solanaceae, Plantaginaceae, and Rosaceae has
identified the pistil S determinant of these families as a stylar-expressed RNase, called
S-RNase (Anderson et al., 1986; Sassa et al., 1992; Ushijima et al., 1998). Sometime back
researchers did site-directed mutagenesis of His93 to Asn93, which turned the Petunia plant
incapable of rejecting incompatible self-pollen due to damaged RNase activity.
During these times when the population is increasing uncontrollably, for feeding people, SI
crops are being preferred as seed setting is not desired. Also, since long breeders have
performed a targeted genetic modification of S genes to successfully generate SI lines of SC
plants. Generation of transgenic SI A. thaliana line by transferring both SCR and SRK genes
from sister plant A. lyrata was done many times back (Nasrallah 2002). Because of its
fruitful aspects in both academia and agriculture, SI has been deeply studied in a diversity of
plant species including some prominent crop plants.
Fig. 1.3: The three major evolutionary transitions in the Plant reproductive system.(
Self-created with reference to Spencer C. H. Barrett et al.; 2010)
3. Conclusion
Exploring the imperative properties of the floral phenotype requires more collection of
genetic data based on morphological, developmental, and evolutionary research. Researchers
have tried to integrate genetics with agriculture giving birth to “modern agriculture”. They
have also been successful in this aspect because of recent technical advancements in genetics
and genomic studies, aiding in the identification of genes and mechanisms important for
producing genetic diversity. The relation between genotype and environment is proved to be
evolutionary, as phenotypic evolution is the result of ecological change through the process of
natural selection. Various influencing genes promoting pollinator-associated floral trait
variation have been identified in several plants. Studies and identification of more candidate
plant S genes will help in targeted modification and development of hybrid crops having
better production and survival traits.
4. Future Perspectives
A huge potential lies in comparative molecular analyses of plants like Thalictrum which can
undergo both wind and animal pollination. Identification of more genes and their
corresponding mutations resulting in floral trait variation concerned with pollinator
specificity is important to understand how pollinator shifts occur between closely related
plant species. Only a few studies have been done with plants of agronomical importance. The
yields of crops with high value such as fruits, nuts, and vegetables depend majorly on
pollination success. Therefore, it the important to take into consideration their floral
metabolic traits while doing plant breeding. GLR(GLUTAMATE RECEPTOR-LIKE
channel) gene is an important candidate gene to reveal the role of Ca2+ signaling in SI
response, further studies are important to understand the exact biochemical mechanism and if
its function is haplotype-dependent. Also, future studies are required to understand the
complete influence of phosphorylation and Ca2+on grass SI response. The mechanism behind
the detoxification of S-RNases in compatible pollen also needs to be further studied. Future
studies should work on the hypothesis regarding the collaborative recognition system and
increase in differentiation of new S specificities in plants. Also, more in-depth studies are
required for a better understanding of the dynamics between landscape and the genetic
diversity of key plant species.
References
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https://doi.org/10.1093/aob/mcp122
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