Molecular mechanism
of sporophytic plants self-incompatibility:
- Sporophytic plants includes most of the land plants and
multicellular algae. The sporophyte represents a diploid
multicellular stage in plants (angiosperms-flowering plants,
gymnosperms-seed plants) or algae exhibiting alternation of
generation or alternation of phases.
- The mechanism of self-incompatibility (SI) is essential, to
prevent self-fertilization or inbreeding. SI also aids in
outbreeding and genetic diversity.
- SI locus genes are highly polymorphic.
- The SI is determined by product of S allele. The S-locus
comprises of multiple (like S1 and S2 locus), tightly linked
genes.
- Two major types of SI exhibited are:
1. Gametophyte SI (GSI):
- Most common form of SI, occurring in most angiosperms, like
Solanaceae, beets, grasses, roses, etc.
- S- allele product is functional after meiosis. SI controlled by
single S-allele.
- Initially both compatible and incompatible pollens germinate
into pollen tubes and begin to grow.
- The fate of the gamete is determined by interaction of pollen
tube and the style.
- The either pathway may result in preventing self or compatible
pollen tube:
a. Gametophyte SI (GSI) based on based on selective
degradation of pollen RNAs- Within the style, S-loci
synthesize, ribonuclease (S- RNAse), which on entering the pollen
tube destroys pollen RNA. This results in halting of pollen
tubes.
b. Gametophyte SI (GSI) based on based on Calcium
dependent signaling cascade (MARK)- signaling cascade
causing pollen cell death.
2. Sporophyte SI (SSI):
- Observed in Asteraceae, Brassica, etc.
- Self-pollen rejection controlled by diploid genotype.
- Three tightly linked gene loci are involved:
- . SLG- S- locus glycoprotein, encodes proteins
(receptors) forming part of stigma cell wall. SLG enables SRK
activity.
b. SRK- S- locus receptor kinase, encodes
transmembrane protein embedded in stigma cell membrane.
c. SCR- S-locus cysteine rich, encodes soluble
ligand similar to that secreted by pollens, binding to same
receptors.
- ARC1- Armadillo repeat 1, degradation of
stigma proteins.
- . MLPK – M-locus protein kinase Positive SI
mediator.
- Incompatibility reaction occur in the stigma papilla cells, by
recognizing self-proteins.
- As observed in S1-S2 system (S1S2 pollen cannot germinate on
S1S3 stigma)
- Pollen will not germinate if the stigma contains either of the
two alleles. S1S2 pollen, produces both SCR1 and SCR2. If either
SCR bind to receptor on the pistil, the kinase activities are
triggered, leads to subsequent events like proteasome destruction
by ubiquitination of proteins.