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Peroxisomes

Aimee Terauchi and Valerie Villareal

History of Peroxisomes
First

observed by electron microscopy in animal cells (1950s), then in plant cells (1960s)

Christian

deDuve (1965)

Isolated from liver cells by centrifugation Called them peroxisomes because they generate and destroy H2O2

The Peroxisome
Single

membrane Roughly spherical

0.2 - 1.7m

Composition

varies

Number and Size of Peroxisomes Vary Depending on Environment

Glucose limited

More glucose limited

Methanol limited

Hansenula polymorpha cells

Protein Import

C-terminal signal sequence: SKL N-terminal signal sequence: RLX5HL Proteins involved in import: peroxins Import driven by ATP hydrolysis Dont have to be unfolded for import

Two Models for Peroxisome Biogenesis

Metabolic Functions of Peroxisomes


Yeasts Biosynthesis: lysine Degradation: amino acids, methanol, -oxidation of fatty acids, decomposition of hydrogen peroxide, glyoxylate cycle Fungi Biosynthesis: penicillin Degradation: -oxidation of fatty acids, decomposition of hydrogen peroxide, glyoxylate cycle Plants Degradation: purines, some reactions of photorespiration (the conversion of glycolate to glycine and of serine to glycerate), -oxidation of fatty acids, decomposition of hydrogen peroxide, glyoxylate cycle Mammals Biosynthesis: ether phospholipids (plasmalogens), cholesterol and bile acids, polyunsaturated fatty acids Degradation: amino acids, purines, prostaglandin, polyamines, -oxidation of fatty acids, -oxidation of fatty acids, decomposition of hydrogen peroxide Humans Biosynthesis: ether phospholipids (plasmalogens), cholesterol and bile acids, polyunsaturated fatty acids Degradation: amino acids, purines, -oxidation of fatty acids, -oxidation of fatty acids, decomposition of hydrogen peroxide

Major Metabolic Functions of the Peroxisome in Plants


1.

2.
3. 4. 5.

-oxidation of fatty acids Glyoxylate cycle Photorespiration (Glycolate pathway) Degradation of purines Decomposition of hydrogen peroxide

Two Types of Peroxisomes in Plants


Leaves

Catalyzes oxidation of side product of CO2 fixation in photorespiration

Germinating

seeds

Converts fatty acid in seed lipids into sugars needed for growth in the young plant

Glyoxysomes

-oxidation occurs in mitochondria and peroxisomes in mammals, but exclusively in the peroxisome in plants and yeast.

Glyoxysomes and Leaf Peroxisomes are Interconverted During Development

Immunogold particles of 2 sizes bound to:

Enzymes of glyoxylate cycle Peroxisomal enzymes

Greening cotelydons

The same population of peroxisomes assumes different metabolic roles depending on developmental stage of cotelydon

Photorespiration and Glycolate


Oxygenase activity of rubisco
Consumption of O2

Glycolate cycle
Production of CO2 Involves 3 organelles (chloroplasts, peroxisomes, & mitochondria)

The Glycolate Cycle

Glycolic acid oxidase

H2O2 production

The Glycolate Cycle

Purine Degradation
Nucleic

acid purine moieties (adenine and guanine) are degraded to uric acid
O2 H2O2 O2 H2O2

xanthine

uric acid
Urate oxidase

allantoin

Xanthine oxidase

Del Rio et al., J. Exper. Botany 2002

Urate Oxidase

High urate oxidase concentrations contribute to formation of crystalline inclusions All purine degradation leads to uric acid
H2O2 O2
O H N C N C H N H N H

H2O2

O2
O H N C

purines

H N H C N

N C H N H

FAD

FADH 2
O

FAD FADH 2
O

H N C O

H2O xanthine oxidase

H2O xanthine oxidase

N H

N H

hypoxanthine

xanthine

uric acid

Oxidases

The oxidases use molecular oxygen to remove hydrogen atoms from specific organic substrates A variety of compounds, including L-amino acids, D-amino acids, polyamines, methanol, urate, xanthine, and very-long-chain fatty acids, serve as substrates for the different oxidases

Peroxide Detoxification
Oxidases use O2 to oxidize organic substances and produce hydrogen peroxide (H2O2) -- e.g., H2O2 generated by glycolate oxidase reaction, -oxidation of fatty acids
Peroxisomes also contain catalase, the enzyme that degrades H2O2.

Importance of H2O2 degradation


2H2O2

2H2O + O2 Peroxisomes contain a high concentration of catalase, a heme protein


catalase
H - - O - - O - -H HO-OH (?)

Other

reactive oxygen species (ROS) are formed in peroxisomes

Reactive Oxygen Species


1eO2 O2 1eH2 O2 1e OH- + OH 1eH2 O

.. . .. . . .. .. .. .. - .. O : H H : O :H : : : : : : : :O : : : + O O O H O O H H O . . . .. .. . .. . . .. ..
Superoxide Hydrogen Hydroxyl anion peroxide radical (radical) Cause damage to lipids, proteins, DNA Amount ROS is reduced by catalase, and superoxide dismutase (SOD) 2O2O2 + H2O2

Radical Chemistry
Initiation: RH + O2 -->R + OH
Termination: R + R --> RR R + ROO--> ROOR ROO + ROO --> ROOR + O2

Propagation: R + O2 --> + ROO

ROO + RH --> R + ROOH


ROOH--> RO + HO

Other Peroxisomal Enzymes

Conclusions

Compartmentalize! To protect the cell from these destructive byproducts, such reactions are segregated.

Peroxisomal Diseases
Adrenoleukodystrophy: Deficiency in oxidation of very longchain fatty acids

Zellweger syndrome: Defect in protein import, giving rise to ghost peroxisomes