Molecular Basis of SI

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University of Horticultural

Sciences, Bagalkot
Presentati
on
On

“Molecular Basis of Self Incompatibility ”

KAVITA PUJARI
UHS17PGD248
Ph.D (Hort.)

College of Horticulture,
Self-Incompatibility
2

Self-Incompatibility was first reported by Koelreuter


in Verbascum phoenicium in 1764

“The inability of a fertile hermaphrodite seed plant


to produce zygote after self-pollination”


(de Nettancourt, 1977)

More than 300 species belonging to 70 families of


angiosperms show self-incompatibility
Self-Incompatibility
3

 Pollen grains are functional but fail to


fertilize egg cell of the same plant.
 Pollen grains fail to germinate on the stigma.
 Pollen grains do germinate, pollen tube fails
to penetrate the stigma.
 Pollen tube enters the style but grows too
slowly to effect fertilization.
 Fertilization is effected, but the embryo
degenerate.
Classification of SI
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Bateman 1952
COMPLEMENTARY SYSTEM
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 Stimulatory type of SI.


 Pistil and pollen which differ in SI

genotype, together provide


substance which stimulate pollen
germination and pollen tube growth.
 But when the genotypes are similar

the germination and growth of pollen


is inhibited.
 Dendrobium
Oppositional system
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 Inhibitory type of SI
 Pollen and pistil produce

substances which prevent pollen


germination if pollen has same SI
reaction as pistil.
 Germination and growth of pollen

differing in SI reaction in not


inhibited.
Self
Incompatibi
7
lity

Homomorp
Heteromorp
hic System
hic System

Gametophy Sporophytic
tic System System

Lewis
Heteromorphic System
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 Flowers of different incompatibility groups are


different in morphology.
 Distyly – single locus ; Tristyly – two loci (S and M)

 Distyly occurs in 24 families and more than 164

genera of flowering plants, the most prominent of


which are the Primulaceae, the Rubiaceae, and the
Plumbaginaceae
 In Primula- 2 types of flowers

Pin(ss) - long style and short stamens


Thrum(Ss) - short style and long stamens
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Dimorphic flowers of buckwheat

Examples: Primula, sweet potato,


Buckwheat, Lythrum salicaria,
Heteromorphic Self
Incompatibility
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Schematic presentation of the intra-morph


incompatibility response in buckwheat
Tristyly
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 Tristylic species are


less widely
distributed, found in
Lythraceae,
Oxalidaceae, and the
monocotyledonous
family
Pontederiaceae
Oxalis alpina
Homomorphic System
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 Incompatibility is not associated with


morphological differences among flowers.
 Found in majority of self incompatible species.
 Incompatibility reaction of pollen is controlled by:
a. the genotype of the plant (Sporophytic control)
b. its own genotype (Gametophytic control)
 2 types:
a. Gametophytic system
b. Sporophytic system
Gametophytic SI Sporophytic SI
 First described by East and  First reported by Hughes and
Mangelsdorf in 1925 in Babcock in Crepis foetida and
Nicotiana sanderae. by Grestel in Parthenium
 Outcome of the interaction argentatum in 1950
between the pollen tube and the  Outcome of the interaction
style is determined by the between the pollen tube and
genotype of the pollen (gamete) the style is determined by the
genotype of the sporophyte
 S-locus products are synthesized
(diploid tissue)
after completion of meiosis
 S-locus products are
 Growth of the pollen tube arrests
synthesized before completion
in the style
of meiosis
 Seen in Solanaceae, Rosaceae,  Growth of the pollen tube
Leguminoseae, arrests at the surface of the
Scrophulariaceae, Papaveraceae, stigma
Onagraceae, Campanulaceae,  Compositae, Cruciferae,
and the monocotyledonous
Convolvulaceae, Betulaceae,
family Commelinaceae 13
Caryophyllaceae and
Sterculiaceae
Sporophytic SI
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S1S2x S1S2 S1S2x S1S2x S1S2x S1S2 S1S2x S1S3 S1S2x S3S4
S1S3 S3S4

S1S2 S1S2 S1S2 S1S2 S1S2 S1S2

Fully Partially Fully Fully


Fully incompatible
incompatibl incompatiblecompatible compatible
e
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MOLECULAR BASIS OF S
Molecular Mechanism Of SI

 3 types:
The Brassicaceae type SI
mechanism

The Papaveraceae type SI


mechanism

The Solanaceae type SI


mechanism
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S Locus
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 Previously S-locus was assumed to be a


single gene but after 1987 molecular
studies revealed S-locus to be much more
complex.
 Using PFGE, it was possible to detect 2
genes within S-locus to control SI.
 The S-locus consists of at least two linked
transcriptional units arranged in pairs.
 One unit function as male determinant
while other as female determinant.
 Highly polymorphic genetic locus.
The Brassicaceae Type SI
Mechanism
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 The incompatible pollen grain is inhibited


at the stigma surface.
 S receptor kinase (SRK) is found in the
plasma membrane of stigmatic cell
(pistil).
 S- locus Glycoproteins enhances the SI
reaction
 S- locus Protein-11/SCR is a small (<10
KDa) protein and predominantly occurs in
anther tapetum.
 Positive effectors are M-Locus Protein
Molecular Mechanism of SI
in Brassicaceae
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SRK get activate when pollen ligand, SP11/SCR, binds to its


extracellular domain.

Dimerization of SRK

Auto-phosphorylation of SRK

Generate cytosolic signal in MOD, an aquaporin

Differential regulation of water- channel protein to inhibit pollen


growth
Model for Molecular Mechanism of SI in
Brassicaceae
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Papilla cell

Takayama et al. 2005


The Papaveraceae type SI
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mechanism
 Inhibition of pollen tube growth occurs at the
stigma surface, before or immediately after
germination
 The stigmatic S proteins are small ( ̴15 kDa)
secreted proteins
 SBP (S protein binding protein) specifically
binds S-proteins
 Interaction of self-pollen with the stigma
induces a calcium-dependent signaling
cascade leading to programmed cell death in
the pollen (Thomas & Franklin-Tong 2004)
Molecular Mechanism of SI in
Papaveraceae
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Binding of the stigmatic S protein to the pollen


S receptor

triggers a signal transduction cascade in the


pollen tube

Interferes the intracellular concentration


gradient of Ca++

Arrests pollen tube elongation and PCD


The Solanaceae type SI
mechanism
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 This type of SI mechanism is found in 3 families:


a) Solanaceae
b) Rosaceae
c) Plantaginaceae
 It is characterized by the arrest of the incompatible

pollen tube within the upper one-third part of the


style.
 S-Locus F box protein was first identified from

Antirrhinum hispanicum as a pollen-expressed, S-


linked gene
 Function of SLF proteins is to promote ubiquitin-

mediated protein degradation.


Molecular Mechanism of SI in Solanaceae
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SLF preferentially interact with its non- self S-
RNase in the cytoplasm of pollen tube

Formation of E3-ubiquitin like complex

Mediates ubiquitination and degradation of non-


self S- RNase by the 26S proteasome

No rRNA degradation inside pollen tube

Pollen tube growth


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 An HT protein and other stylar glycoproteins


have been seen to interact with S-RNases.
 HT, a gene from Nicotiana alata, which encodes
a small asparagine-rich protein of unknown
function, is essential for the SI response in
Nicotiana and Lycopersicon.
 It is proposed that S-RNases bind these
components in vivo, and that these complexes
might enable the uptake of the S- RNases into
the pollen tube. This interaction is probably
required to achieve pollen tube inhibition.
McClure, B.A. et al.
(2000)
Model for Molecular Mechanism of SI
in Solanaceae
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Takayama et al. 2005


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1) Single Cross

2) Double Cross

3) Three line cross


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Line A Line B
S11 S22

S11 X S22

( A X B)
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 Simplest method.
 Method for developing single cross:

a) Two SI, but cross compatible, lines are


inter planted, seed obtained from both
the lines would be hybrid seed.
b) A SI line may be inter planted with a self
compatible line and seeds from only the
SI line would be hybrid.
 Hybrids are uniform one.
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 Firstly utilized by Thompson (1959) for


the production of Maris Kestrel, a
marrow- stem kale variety.
 Requires more preparatory breeding work
like isolation of four SI lines and testing
for combining ability.
 Hybrid is less uniform.
 Yield lower than single- cross hybrid.
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Line Line B Line C Line D


A
S11 S22 S33 S44

S11 X S22 S33 X S44

Ax CX
B X D

{( A X B) X ( C X D)}
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 Suggested to reduce the cost of


producing inbred parental lines.

 Final hybrid is far from uniform, may be


regarded as a synthetic variety.

 Problematic to produce and maintain


parents
Three line Cross Hybrid

X
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Inbred X Inbred Inbred Inbred
A B D E

Ax X Inbred C Dx X Inbred
B E F

X S46, S56
S13,
ABC
S23 DEF

TCH
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 New incompatibility reactions.


 Depression by Continuous Inbreeding
 High cost of hybrid seeds
 Reduction of Incompatibility by
Environmental conditions
 Restriction of pollination within
Parental lines
35  SI was one of the most important pollination systems for hybrid
seed production in B. napus
 It was once the only system used in B. napus hybrid production
in China prior to the discovery and use of the cytoplasmic male
sterility (CMS) system (Fu 1981).
 When compared to CMS-
a) SI hybrids have higher F1 seed production,
b) No obvious negative cytoplasm effect, and
c) an abundance of restorers.
d) It was found that F1 hybrids produced with SI lines showed
heterosis in B. napus.
 SI hybrids-
a) Youyan 3 (China, 1991),
b) HC-120 (Canada, 1991),
c) Huayuyou 640 (China, 2009), and
d) Huayouza 95 (China, 2010).
Contd…
36

 The other hybrid BR CMS × FT showed


to be less uniform, forming smaller
and lighter curds.
 During Seed production-
 Self-incompatibility appears to be
more effective than
cytoplasmic male sterility.

But CMS system provides much more


reliable sterility than self-incompatibility.
37

Crops hybrids
Cauliflower Pusa Hybrid-2, Snow Queen,
Snow King, White Contessa

Cabbage BRH-5, H-44, H-43, Pusa


Synthetic, Meenakshi

Chinese Hamburg-3
Cabbage
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 The F1 hybrid of SI line Montano × self-pollinating


line FT 13 turned out to be the best combination
in a preliminary field trial.
 Good uniformity,
 High curd quality,
 good covering of curd by inner leaves
 A satisfactory disease resistance was also seen
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Curd of the hybrid of SI line Montano × self-


pollinating line Fortuna FT 13
Hort. Sci. (Prague), 33, 2006 (4): 148–152
Self-Incompatible genotypes used as variety-

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Crops Genotypes
Cauliflower IIVR – 1, IIVR 50, Kataki Early- 29,
HAZIPUR 4 (BP), Pusi 4, Agahani JBT-
23/60 , late Agahani, Agahani-8, Pusa
Hazipur, Agahani Long Leaf, Agahani
Small Leaf, kuwari 1, Kataki-12
Radish Pusa Chetaki, Pusa Desi, Half Red, Acc
No. 30205, Acc No. 282, Chinese Pink,
BDJ 689, Desi Red, Khasi Kata
CONCLSION
41

 In recent years, exciting progress has


been made towards understanding the
molecular basis of several self
incompatibility systems. However, still
there is much to be discovered about
processes involved in this system.
 It is hoped that current research in self
incompatibility mechanisms may come
out with a simplified treatment which has
a sound practical utility.
42

Thank You

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