Cytoplasmic Inheritance

Joe Ironside
 Mendelian inheritance
• Two copies of each
gene are inherited
– One from each parent
• Females
– All chromosomal genes
• Males
– All chromosomal genes
apart from those on the
X and Y chromosomes
 Non-Mendelian inheritance
Y chromosome X chromosome of
of males males
• Only one copy of a
gene is inherited
– From the male parent
• Y-linked genes of males
– From the Female
parent
• X-linked genes of males Cytoplasmic genes

• Cytoplasmic genetic
elements
 Question
• Which genes are inherited through the
cytoplasm?
 Cytoplasmic genes
• Organelle genomes
– Mitochondria
• Hydrogenosomes
– Plastids
– Nucleomorphs
• Bacterial endosymbionts
– Primary endosymbionts
– Secondary endosymbionts
– Parasites
• Eukaryotic endosymbionts
– Parasites
Organelle genomes
 Mitochondrial symbiosis
• Occurred once
– All mitochondria share a
mitochondrial common
ancestor
• Occurred at or shortly
after origin of eukaryotes
– All extant eukaryotes
• possess mitochondria
– or are descended from
ancestors which possessed
mitochondria
 Plastid symbiosis
• Occurred once
– All plastids share a
common ancestor that
was also a plastid
• Occurred after
eukaryotes had
diversified
– Probably in a
biflagellate protist
 Secondary endosymbiosis
• Non-photosynthetic
eukaryote
– Engulfs photosynthetic
eukaryote
• Produces eukaryotic
“chloroplast”
– With four membranes
» Host endomembrane +
plasma membrane + 2
original chloroplast
membranes
– May have vestigial nucleus
 Spread of plastids
• Secondary and
tertiary
endosymbiosis
– Spread plastids
between different
eukaryote lineages

Keeling (2010) Philos Trans R Soc Lond B Biol Sci. 365: 729–748
 Uniparental
inheritance
• Animals and plants
– Organelles are excluded
from sperm or pollen
• Cytoplasmic genes are
transmitted only though
females + -
• Chlamydomonas
rheinhardtii
– Parents are of + and –
mating types
• Organelles of – parent are
destroyed after cell fusion
 Bacterial endosymbionts
• Primary endosymbionts
– Present in all individuals
within a population (but
not all cells)
• E.g. Buchnera in aphids
• Secondary
endosymbionts
– Present in some
individuals within a
population
• Parasites
– Present in some
individuals within a
population
• Negative effect on fitness
– E.g. Wolbachia
 Eukaryotic endosymbionts

• Parasites
– Microsporidia
– Paramyxids
 Reduction of endosymbiont
genomes
• Inefficient purifying selection
– Small population size
– No opportunities for
recombination
– Muller’s Ratchet
– Selective sweeps
– Background selection
• Sheltered environment
• Gene transfer
– To host nucleus
• Results
– Loss of genes
– Amplification of genes for
amino-acid synthesis
Reduction of organelle genomes
• Mitochondrial
genome
– Reduced size
• Gene loss
– Metabolites
imported from
cytoplasm
• Transfer of genes
to the nucleus
– Proteins imported
from cytoplasm
 Horizontal
gene transfer
• Organelles
– encode only 1-5% as
many proteins as free-
living prokaryotes
• Organelle proteins
– encoded by nuclear genes
– Imported to organelles
• Energy metabolism
• Amino acid synthesis
• Nuclear genes
– derived from prokaryotic
genes
• Implies gene transfer
from organelle to nucleus
 Strange and degenerate
mitochondria
• Hydrogenosomes
– Derived from mitochondria
– Produce ATP anaerobically
• Use proteins acquired from anaerobic
bacteria and located in the nucleus
– Lack genomes
• (with the exception of Nyctotherus
ovalis, a parasite of the cockroach
gut)
• Mitosomes
– Relict mitochondria
• Lack genomes and do not produce
ATP
 Organelle proteomes
• Less than 20% mitochondrial proteins
derived from original symbiont
 Question
• How could cytoplasmic genes increase
their transmission?
 Strategies to increase
cytoplasmic transmission
• Increase fitness of female offspring

X
– Enhance metabolism
– Reduce sibling rivalry by killing males
• Increase proportion of female
offspring
– Convert males or hermaphrodites into
females
• Kill offspring of unrelated females
– Cytoplasmic incompatibility

X
Metabolic enhancement
• Mitochondria
– ATP synthesis
• Plastids
– Photosynthesis
• Aphid endosymbionts
– Amino acid synthesis